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hHr 3 1387 

L16I— O-1096 









Published by 


MAY 28, 1965 

101 B'JRRlLLHAli- 


A Continuation of the 














Curator, Division of Mammals 



The Late Curator of Mammals, 
American Museum of Natural History 



Published by 


MAY 28, 1965 

Edited by Edward G. Nash 

Library of Congress Catalog Card Number: 65-22^9 




List of Text Figures 5 

Introduction 7 

Acknowledgements 8 

Scope 10 

Material 11 

Methods 13 

Color 14 

Form of Presentation 15 

Size and Dimensions 15 

Order 16 

Taxonomic History 18 

Classification 20 

Giant Squirrels, Genus Ratufa 29 

Species macroura, Ceylonese Giant Squirrel 34 

Species indica, Indian Giant Squirrel 41 

Species bicolor. Pied Giant Squirrel 49 

Indian striped Squirrels, Genus Funambulus 65 

Species pennanti, Five-striped Indian Squirrel 72 

Species palmarum, Indian Palm Squirrel 77 

Species tristriatus, Western Ghats Squirrel 87 

Species layardi, Sinhalese Jungle Squirrel 91 

Species sublineattis, Dusky Indian Jungle Squirrel 95 

Common Tree Squirrels, Genus Callosciurus 99 

Species erythraeus, Red-bellied Squirrel 101 

Species flavimanus, Belly-banded Squirrel 120 

Species /errM^mcMS, Pegu Red Squirrel 158 

Species /fn/ai/som, Siamese Squirrel 161 

Species caniceps, Tenasserim Squirrel 187 

Species phayrei, Shan Highlands Squirrel 201 

Species inornatus, Laotian Squirrel 209 

Species pygerythrus, Irrawaddy Squirrel 213 

Striped Tree Squirrels, Genus Tamiops 231 

Species mcclellandi, Burmese Striped Tree Squirrel 235 

Species rodolphei, Cambodian Striped Tree Squirrel 243 




Species smnhoei, Chinese Striped Tree Squirrel 246 

Species maritimus, Maritime Striped Tree Squirrel 251 

Plain Long-Nosed squirrels, Genus Dremomys 259 

Species lokriah, Himalayan Long-nosed Squirrel 263 

Species pernyi, Chinese Long-nosed Squirrel 269 

Species rufigenis, Red-cheeked Long-nosed Squirrel 278 

Species pyrrhomerus, Red-hipped, Long-nosed Squirrel 283 

Species evereiti, Montane Bornean Long-nosed Squirrel 289 

Indochinese Ground Squirrel, Genus Menetes 293 

Species berdmorei, Indochinese Ground Squirrel 294 

Chinese Rock Squirrels, Genus Sciurotamias 303 

Species davidianus, Chinese Rock Squirrel 304 

Species forresti, Chinese Cliff Squirrel 309 

Summary 311 

Gazetteer 313 

References 337 

Index 345 



1. Skull of pied giant squirrel, Ratufa bicolor 30 

2. Range of Sinhalese giant squirrel, Ratufa macroura 35 

3. Range of Indian giant squirrel, Ratufa indica 40 

4. Range of pied giant squirrel, Ratiifa bicolor 50 

5. Skull of Western Ghats squirrel, Funambulus tristriatus 67 

6. Skull of Indian jungle squirrel, Funambulus sublineatus 70 

7. Range of five-striped Indian squirrel, Funambulus pennanti 73 

8. Range of Indian palm squirrel, Funambulus palmarum 78 

9. Range of Western Ghats squirrel, Funambulus tristriatus 88 

10. Range of bright-striped squirrel, Funambulus layardi 92 

11. Range of Indian jungle squirrel, Funambulus sublineatus 96 

12. Skull of Irrawaddy squirrel, Callosciurus pygerythrus 101 

13. Ranges of red-bellied squirrel, Callosciurus erythraeus, and the belly- 
banded squirrel, Callosciurus flavimanus 122 

14. Ranges of Pegu red squirrel, Callosciurus ferrugineus, and Thailand tree 
squirrel, Callosciurus finlaysoni 162 

15. Ranges of the Tenasserim squirrel, Callosciurus caniceps; Shan Highlands 
squirrel, C. phayrei; and Laotian squirrel, C inornaius 203 

16. Range of the Irrawaddy squirrel, Callosciurus pygerythrus 214 

17. Skull of striped tree squirrel, Tamiops svnnhoei 233 

18. Ranges of striped tree squirrels, Tamiops mcclellandi and T. rodolphei . . 236 

19. Ranges of striped tree squirrels, Tamiops sunnhoei and T. maritimus (part) 247 

20. Range of striped tree squirrel, Tamiops maritimus (part) 252 

21. Composite ranges of species of the genus Dremomys 260 

22. Skull of Himalayan long-nosed squirrel, Dremomys lokriah 261 

23. Skull of Chinese long-nosed squirrel, Dremomys pernyi 261 

24. Range of Himalayan long-nosed squirrel, Dremomys lokriah 264 

25. Range of Chinese long-nosed squirrel, Dremomys pernyi 270 

26. Range of red-cheeked squirrel, Dremomys rufigenis 279 

27. Range of red-hipped squirrel, Dremomys pyrrhom^rus 284 

28. Orbitonasal length histogram for species of Dremomys 285 

29. Length of orbit histogram for species of Dremomys 286 

30. Skull of Siamese ground squirrel, Menetes berdmorei 293 

31. Range of Siamese ground squirrel, Menetes berdmorei 295 

32. Skull of Chinese rock squirrel, Sciurotamias damdianus 304 

33. Ranges of Chinese rock squirrel, Sciurotamias damdianus, and Chinese 
cliff squirrel, S. forresti • 305 



The present study was initiated in connection with the identifi- 
cation of a large collection of Oriental squirrels acquired by the 
Archbold Expeditions of the American Museum of Natural History. 
Contemplating a taxonomic review of the Oriental squirrels, Dr. 
George Tate undertook to re-examine and attempt to identify with 
the population to which it belongs, each of the approximately 700 ac- 
cessible type specimens of squirrels from southeastern Asia and the 
East Indies. This study involved visiting many of the museums in 
which the types are deposited, and these visits included study of 
extensive additional collections of Oriental squirrels. Procedure con- 
sisted of vocally recording new observations on the types in Ameri- 
can and European museums, of photographing the skulls of types, 
and of making detailed measurements of them. Comparative obser- 
vations on other specimens of Oriental squirrels in European muse- 
ums in 1951 were also recorded. In some instances specimens were 
sent on loan for examination (by Tate). 

Dr. George H. H. Tate died in December of 1953. The other 
author (Moore) was invited to put the manuscript on the Oriental 
squirrels into final form for publication. Gradually he found that 
the manuscript treatment of many genera could not be put into form 
for publication by editing. He then undertook to change the concept 
of the paper from a review to a substantial revision based on Tate's 
transcribed observations of European material and new study of 
about 8000 Oriental squirrel specimens in museums in the United 
States. Beginning in September, 1957, grants were soHcited from 
federal agencies, but not for the preparation of a taxonomic revision 
alone. It was pointed out that this is a study of a subfamily of ter- 
restrial mammals unusually variable in color, in a geographic area 
of exceedingly varied land form and extraordinarily active orogeny 
and erosion. Federal support of the morphological study and revi- 
sion was sought and obtained on the grounds that they will reveal 
evidence on the evolution of the animals in relation to land forms and 
water barriers and thus may be important to evolutionary theory. 

Discoveries of special taxonomic importance in this revisionary 
work have led to several separately published contributions. One is 


a classification of the diurnal squirrels of the world (Moore, 1959). 
Another is an assessment of the taxonomic importance of the num- 
ber of pairs of functional mammae in the Sciurinae and also the 
relationship of this character in Sciurinae to climate and to size of 
broods (Moore, 1961a). The strengthening of the classification of 
the living Sciurinae provided in these two papers invited considera- 
tion of the evolutionary implications of the geographic distribution 
of the morphological characters observed. These implications have 
been treated in part in the 1959 paper that offered the classification, 
in part in a paper on the distribution of squirrels in the Indian Sub- 
region (Moore, 1960), and further in a discussion of spreadings across 
the Bering and Panamanian land bridges (Moore, 1961b). 

The use of the plural pronoun "we" should not confuse the reader 
who has read the above chronology of authorship. The "we" is 
usually meant only to imply participation of both authors in obtain- 
ing and collating the information and is not intended to place final 
responsibility for decisions on more than one pair of shoulders. Vir- 
tually no decision or opinion stated is Tate's alone unless so indicated 
in text except for ones that could only have been his because they 
could have been reached only by a study of the material in European 


One or both of the authors are indebted to the following persons 
in the United States who have welcomed us to their offices and pro- 
vided us access to the study collections in their charge: Dr. David H. 
Johnson, United States National Museum, Washington, D.C.; the 
late Colin C. Sanborn and Mr. Philip Hershkovitz, Chicago Natural 
History Museum, Chicago, Illinois; Miss Barbara Lawrence, Mu- 
seum of Comparative Zoology, Cambridge, Massachusetts; Dr. H. 
Radclyffe Roberts (and Dr. Karl F. Koopman, formerly of) Acad- 
emy of Natural Sciences, Philadelphia, Pennsylvania; and Dr. W. H. 
Burt and Dr. Emmett T. Hooper, Museum of Zoology, University 
of Michigan, Ann Arbor. 

In Europe the following persons extended similar courtesies and 
privileges to Tate or helped us with loans and information, for 
which we both express our appreciation here: Mr. T. C. S. Mor- 
rison-Scott and Mr. R. W. Hayman, both of the British Museum 
(Natural History), London; M. Jean Dorst, Museum National d'His- 
toire Naturelle, Paris; Count Nils Gyldenstolpe, Royal Natural 
History Museum, Stockholm; Dr. L. Forcart, Naturhistorisches 


Museum, Basel; Dr. A. M. Husson, Rijksmuseum van Natuurlijke 
Historie, Leiden; Dr. E. Stresemann, Mr. K. Zimmermann, and 
Dr. W. Meise, Zoologisches Museum der Humboldt Universitat, 
Berlin; Dr. R. Mertens, Senckenbergisches Naturforschende Gesell- 
schaft, Frankfurt-am -Main; Dr. C. Stop-Bowitz, Universitetets Zoo- 
logiske Museum, Oslo; Dr. A. Holm, Zoologiska Institutionen, 
Uppsala; and the late Mr. H. J. V. Sody, of Amsterdam. 

For loans of important material from museums in the Oriental 
Region we are grateful to the late Dr. Sunder Lai Hora and to Dr. 
Biswamoy Biswas, both of the Zoological Survey of India; to Mr. 
Humayun Abdulali, of the Bombay Natural History Society; and 
to Mr. P. E. P. Deraniyagala, of the National Museums of Ceylon. 

This investigation was supported from September, 1957, through 
August, 1961, in part by Public Health Service research grant 5327 
and in part by National Science Foundation grants 4447, 6250, and 
14265. The surviving author is very appreciative of these. 

Gratitude is expressed to Dr. Harold E. Anthony of the American 
Museum of Natural History for the original invitation (to Moore) 
to take up where Tate had left off, and for support and encourage- 
ment until his retirement in May, 1958. His successor, Dr. R. G. 
Van Gelder, continued to make research facilities available for this 
project through the summer of 1961, provided special support from 
departmental funds during one four-month period, and through crit- 
icism of the manuscript has contributed toward accuracy, consist- 
ency, and clarity of expression in this paper. 

Acknowledgment for a special kind of helpfulness is made to 
Mr. Herbert G. Deignan, Curator of Birds, United States National 
Museum, for a great deal of advice and help in the finding and spell- 
ing of place names in Thailand. 

Mr. Melvin A. Traylor, Chicago Natural History Museum, pro- 
vided me with data on the collection localities of Floyd T. Smith in 
Szechwan, China. Dr. Charles Vaurie, American Museum of Nat- 
ural History, welcomed us to study his personal set of 1:1 million 
topographic maps of the Orient. 

The skull drawings are from the pen of Mrs. Richard G. Zweifel. 
The inking and lettering of fourteen of the maps are some of the last 
work of Mrs. D. F. Levett Bradley, who passed away before com- 
pleting the series. The maps showing the distribution of Sciurotamias 
and of Ratufa macroura, R. indica, and of the several species of Callo- 
sciurus were inked and lettered by Mrs. Zweifel. 

10 fieldiana: zoology, volume 48 


The number of tribes of diurnal squirrels represented in the Ori- 
ental Region (5) exceeds that of any other faunal region of the world. 
The number of genera of diurnal squirrels native to the Oriental 
Region (17) greatly exceeds that of other faunal regions (see Table 1). 
Several of the genera listed in Table 1 as Oriental occupy areas mar- 
ginal to the Oriental Region proper. Prosciurillus, Rubrisciurus, and 
Hyosciurus are endemic to the island of Celebes, which lies in an area 
transitional between the Oriental Region and the Australian Region. 
Sciurotamias occupies an area of China which is almost entirely tran- 
sitional between the Oriental and Palearctic regions. Nevertheless, 
none of the 17 genera here listed as Oriental extends more than mar- 
ginally into another of the faunal regions of the world. These data 
suggest that the Oriental fauna of diurnal squirrels may be the most 
important in the world to study. They are also a convenient unit 
to revise. 

The Oriental Region is sometimes divided into three subregions: 
the Indian, the Indochinese, and the Malaysian. In the opinions of 
some authorities the Philippine Islands constitute a fourth Oriental 
subregion, but others consider it transitional between the Orien- 
tal and Australian regions. The diurnal squirrel fauna of the 
Oriental Region is well divided by the subregional boundaries. One 
of us has discussed in detail how the boundary between the Indian 
and Indochinese subregions separates the giant squirrels as species 
and all other diurnal squirrels as tribes (Moore, 1960). The bound- 
ary between the Indochinese and Malaysian subregions provides a 
weaker but still very strong division. This boundary is generally 
(and here) drawn at the constriction of the Malay Peninsula called 
the Isthmus of Kra. (See the discussion by Chasen, 1940, p. x, and 
the map, fig. 13 in the present paper.) Figures 7 and 8 combined 
may be said to indicate crudely the extent of the Indian Subregion 
by distribution of the squirrels. Figure 13 does the same for the 
Indochinese Subregion. The scope of the present paper is revision 
of the Sciurinae of the Indian and Indochinese subregions. A large 
part of it constitutes the first revision of the species of Callosciurus 
of the Indochinese Subregion. 

The gliding squirrels, Petauristinae, which with the Sciurinae con- 
stitute the family Sciuridae, are nocturnal and thus more difficult 
for mammal collectors to obtain. They are rarely collected by orni- 
thologists. Consequently, in American collections of Oriental squir- 
rels, specimens of Petauristinae number about one to every 15 of 


Sciurinae. The Petauristinae, although included in the original plans 
and draft by Tate, have not yet been studied by Moore, but see 
McKenna (1963). 

The geographic area included in the present study is thus pre- 
cisely that part of the Oriental Region included by EUerman and 
Morrison-Scott (1951). For this reason the report here on the giant 
squirrel species, Ratufa hicolor, is confined to the area north of the 
Isthmus of Kra, although the species range extends southeastward 
through Malaya, Sumatra, Java, and Bali. There are, however, five 
other polytypic species with ranges that are widespread in the Indo- 
chinese Subregion but that extend beyond its border down the Malay 
Peninsula for varying distances. These five species are treated in 
full because their ranges do not extend beyond the length of the 
peninsula, and because they generally have but one subspecies south 
of the Isthmus of Kra. 

Four-fifths of the species of the genus Dremomys are widespread, 
polytypic species of the Indochinese Subregion. There is, however, 
a poorly known, apparently monotypic species, everetti, confined to 
Borneo. In the interest of having the revision of the genus Dre- 
momys in one publication, a treatment of everetti is included here. 
The forms from south of the Isthmus of Kra that are treated in the 
present work are labeled "extraterritorial." 


In the lists of material examined for each form, the locality near 
which a specimen was taken is ordinarily given as written by the 
collector on the specimen label. This rule has been departed from 
occasionally when evidence clearly shows that the collector was spell- 
ing the name of one locality in different ways. Since for some Ori- 
ental countries one may find the name of a single locality spelled 
(transliterated to English or French) several different ways on as 
many maps (e.g., Pak Jong, Pak Chong, Pak Xong), the locality 
as spelled by the collector may be followed in brackets by the name 
one of us found on a map and presumed to be the locality meant. 
Often the collector has recorded on the specimen tag two or more 
locality names, indicating a village, the town to which it is nearest, 
and perhaps the provincial capital to which they are both near. If 
it has not been possible to locate the village on a map, the town or 
occasionally even the provincial capital has sometimes been plotted 
as the locality for this specimen on the appropriate species distribu- 



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tion map. The instances in which this has been done are clearly 
indicated as follows: the localities not found (as well as any others, 
such as "India," not suitable for plotting) are placed in quotation 
marks in the list of material examined, and the nearby places that 
are plotted instead are italicized (e.g., the village, subdistrict town, 
district town, and provincial capital "Ban Maeo, Goksatawn," Dan 
Sai, Loei, Thailand). 

In our lists of material examined, the institutions in which the 
study material is housed are cited by initials as follows: 

AMNH — American Museum of Natural History, New York 
ANSP — Academy of Natural Sciences, Philadelphia 
ASB — Asiatic Society of Bengal, Calcutta 
BM — British Museum (Natural History), London 
CBK — Private collection of C. Boden Kloss (not seen) 
CNHM — Chicago Natiu-al History Museum, Chicago 
IM — Indian Museum, Calcutta 
MCZ — Museum of Comparative Zoology, Cambridge, Massachusetts 
MNCN — Museo Nacional de Ciencias Naturales, Madrid 
MNHN — Museum National d'Histoire Naturelle, Paris 
MNHUI — Museum of Natural History, University of Illinois, Urbana 
NM — Naturhistorisches Museum, Basel 
NMC — National Museums of Ceylon, Colombo 

NR — Naturhistoriska Riksmuseum, Stockholm 
RNH — Rijksmuseum van Natuurlijke Histoire, Leiden 
SMT — Staatliches Museum fiir Tierkunde, Dresden 

SNG — Senckenbergisches Naturforschende Gesellschaft, Frankfurt-am-Main 
UMMZ University of Michigan Museum of Zoology, Ann Arbor 
UZM — Universitetets Zoologiske Museum, Oslo 
ZI — Zoologiska Insitutionen, Uppsala 
ZMHU — Zoological Museum of Humboldt University, Berlin 


In many species the present work has been based on remarkably 
fine and extensive collections. Some mainland subspecies are known 
from at least one large series of specimens from a single locality, and 
smaller numbers from a dozen other widely distributed localities, 
providing, thus, some indication of individual variation with which 
to evaluate evidence of geographic variation. Other subspecies are 
represented by much less material, and future collections may en- 
able other students to define better the characteristics and ranges 
of many subspecies. In order to facilitate advancement of knowl- 
edge of these species, we have attempted to present our knowledge 
in detail specific enough to enable future students to add their evi- 
dence to ours, perhaps without the necessity of restudy of the perti- 
nent material on which we report. 


Color. — One of us (Moore) has employed Ridgway's (1912) no- 
menclature, and where capitalized color names are used in this paper, 
direct comparison of the pelage (s) with Ridgway's colors is implied. 
To determine color of a series of study skins, we arranged them on a 
horizontal plane before a window, preferably at a time between 
10 A.M. and 4 P.M. and when the sky was blue, but never when the 
sun could shine directly on the specimens. The skins were oriented 
with tails toward the window, and the color comparison was made 
from a view perpendicular to the surface upon which the specimens 
lay, and thus from a point directly above the part being compared. 

Color comparisons were virtually all made at the institution in 
which the study specimens were stored, but at the United States 
National Museum, because of cloudy weather and distance to win- 
dows, some color readings were taken under direct light of an ordi- 
nary 100-watt, electric light bulb with a tungsten filament. 

In order to avoid the risk of losing or damaging a copy of Ridg- 
way by traveling with it, we borrowed and used different copies at 
the different institutions. Comparisons of material at one museum 
with material at another by Ridgway color names risks variation 
between copies of the book, and is thus relatively seldom employed 
here. Persons seeking to make identifications of considerable im- 
portance are advised in no case to depend solely upon distinguishing 
a small difference in colors in Ridgway (1912), but to send skins and 
skulls to one of the museums listed as having materials of the pre- 
sumed form for confirmation of the identification. It should perhaps 
be pointed out that diagnosis of a subspecies of Oriental diurnal 
squirrel rarely depends upon color of a single part of the pelage which 
is only one Ridgway color chip different from an adjacent subspecies. 

The term "agouti" as used here means body pelage with one or 
more minute contrasting color bands on the individual hairs that, 
in a broad view of the pelage, produce a speckled effect. Except 
where otherwise stated, color of agouti pelage is recorded here as the 
general effect of the visible portions of the pelage, its contrasted ele- 
ments blended by viewing without magnification with the viewer's 
eyes deliberately out of focus. The banding of agouti body pelage 
generally consists of one, two or three light-colored bands (parts of 
the length of a hair sharply differentiated by color) on any particular 
hair, but some hairs in agouti pelage have no bands at all. On the 
individual hairs of the tail, it is the black or blackish bands that are 
counted unless specifically otherwise stated, and the count of black- 
ish bands on individual hairs does not include the almost ubiquitously 


black tip because this tip is often markedly thinner than the remain- 
der of the hair, and often shorter than the blackish bands. 

Form of presentation. — Decision to change the study of the Ori- 
ental squirrels from a review to a revision derived in part from a 
growing curiosity about the origins of the species and from discovery 
of indications that better study would reveal firmer evidence on the 
manner of their evolution. In the Indochinese Subregion, for ex- 
ample, the exceedingly rich and rather well-collected diurnal squirrel 
fauna has been evolving on an orogenically and tectonically active 
mainland that has been sliced into parallel north-south strips of land 
by large rivers that flow down from the Tibetan Plateau. 

Thus the intention has not been primarily the laudable desire to 
bring badly needed order to a chaotic fauna and to complete a con- 
venient revision by means of which virtually all new specimens might 
be identified to subspecies. The emphasis has rather been to revise 
the species for evidence about the evolution of the species from coin- 
cidence of morphological and geographic relationships. Where the 
relationships have been obvious enough from the material examined, 
there has been some tendency simply to plot the subspecies distri- 
bution on the map, record the material examined, and move on: 
Where differences have been more difficult to see, as in the subspecies 
of pygerythrus or the relationships of phayrei, elaborate written analy- 
ses became necessary and have, of course, been incorporated into the 
revision. New evidence on relationships between species has been 
especially sought, and where found is given in detail. 

To synonymize subspecies without explanation is certainly not a 
policy which we would wish to advocate. Generally, where there 
has seemed to be basis enough for a finer splitter to recognize a sub- 
species that we have lumped with another, we have pointed that out. 
But the 6}/2 years spent has been too short, the digressions to exploit 
important discoveries too long, and the pressure from supporters to 
complete a revisionary manuscript too great, to allow the explana- 
tion of why each named form that we have synonymized is by our 
observations undeserving of recognition. 

Size and dimensions. — Students who have studied species that 
are more conservative in pelage color and more variable in skull 
dimensions than diurnal tree squirrels may be surprised at how sel- 
dom we have needed to utilize measurements. Tree squirrels are, 
however, well known to be especially conservative in variation in 
dimensions of the skull. (See Moore, 1959, p. 192 and Table 3.) 
On hearing that the discovery of a skull character that seemed to 


sort tree squirrels into tribes was tempting us to digress, one senior 
mammalogist assured us almost bitterly, "If you discover one good 
skull character in tree squirrels, it will be worth every dollar the 
foundation puts into your project." A number of good skull char- 
acters at the generic level and above were discovered and reported 
(Moore, 1959), but at the species and subspecies level in tree squirrels 
it has been obvious that the return for effort expended would be im- 
practical for a revision so extensive. In the semi-terrestrial long- 
nosed squirrels, however, species differences in skulls could be de- 
tected and are utilized. 

Scientists who have presented, and to an extent utilized, body 
and skull dimensions in their revisions of much smaller taxonomic 
units may find it difficult to endure our failure to emulate their ad- 
mirable intensive methods in our work. Tate obtained and recorded 
body and skull measurements and other data for each type specimen. 
Those of the body he apparently copied from specimen tags, adding 
plus-or-minus to measurements whose accuracy he doubted. He 
measured the skulls himself. Some of these data for each genus are 
presented in Tables 2-4, 6, 8-13, and 15. In these tables total length 
of skull is the greatest occipitonasal length. Mastoid breadth is the 
greatest span of the mastoid processes. Nasal length is the greatest 
length of the longer nasal bone. Diastema is the least distance from 
the alveolus of the incisor to the alveolus of the fourth upper pre- 
molar. Length of palate is the distance from the gnathion to the 
most anterior point on the posterior margin of the palate. Length 
of bulla is the greatest external length of the auditory bulla. Maxil- 
lary toothrow is the crown length of the upper row of cheekteeth 
from the most anterior extension of the crown of the fourth pre- 
molar to the most posterior point on the crown of the third molar. 
Orbitonasal length (used in fig. 28) is taken from the most anterior 
point in the margin of the right orbit to the most anterior point on 
the right nasal. Length of orbit (used in fig. 29) is taken from the 
most anterior point on the margin of the orbit (a small notch in the 
posterior margin of the lacrimal) to the most posterior point on the 
margin of the orbi to- temporal fossa. 

Order. — The order of genera progresses from those with no trans- 
verse septum in the auditory bulla to three such septa, as in the latest 
classification of the Sciurinae (Moore, 1959). This progression is 
considered to be from unspecialized to rather highly specialized in 
this particular character, but it does not imply that Ratufa is unspe- 
cialized in general adaptation to a niche (which would be untrue) or 


that Sciurotamias is highly speciaHzed in general adaptation to a 
niche (though this may be true) . The order of species within a genus, 
and subspecies within a species, is geographical, from a convenient 
edge of the range toward the far edges. 

Under accounts of the genera we give the accepted name, syno- 
nyms if any, type species, definition, and diagnosis. Where it has 
seemed especially valuable, we have included data on systematic 
history and intra-generic relationships, and in several instances have 
provided keys to identification of the species. 

In the account of each polytypic species we have given the ac- 
cepted name, definition, and diagnosis. If it belongs in a polytypic 
genus we have generally commented on its relationships to other 
congeneric species with which it is in geographic contact, or which it 
particularly resembles. Sometimes the species account has seemed 
the proper place for general comments on the intraspecific variation. 
In a few instances a systematic history of the species is provided, 
and in the case of the extraterritorial species Dremomys everetti ob- 
servation of the rarely seen deciduous upper third premolars stimu- 
lated some further investigation and reporting of aspects not treated 
here in the other species. 

In each account of subspecies and monotypic species, we have 
given the accepted name, synonyms if any, information on the type 
specimen (s), and a list of the material examined. If some informa- 
tion regarding the type is lacking from our account, this may be 
taken to imply that we did not find that information with the origi- 
nal materials examined or in the original published account. If our 
data and observations and opinions on a type (generally Tate's, of 
course) differ from any given in the original published description 
of the form, this may be assumed to mean that our information, 
observation, or opinion (recorded by Tate) differs from that of the 
original describer. The list of material examined is generally fol- 
lowed by one or more descriptions. These descriptions may be of 
three kinds, and the kind is indicated by the heading given to each. 
"Original Description" designates a quotation from the original pub- 
lished description, usually of the type specimen, of course. "Type 
Description" designates observations by one of us of the type speci- 
men or the cotypes. Unless dated after 1953, these are Tate's ob- 
servations, expanded from his telegraphic notes. "Pelage Color" 
designates descriptions from series of specimens considered to repre- 
sent populations constituting the subspecies under consideration. 
These are given in Ridgway's color terms, and the observations were 


made in the manner described above in the account of color. Dis- 
cussion of some especially interesting or significant features is some- 
times contributed under the heading "Variation," "Dimensions," or 
"Discussion," and field observations from the literature are occasion- 
ally quoted under the heading "Habits." When only one small 
paragraph of comment follows the list of material examined, no head- 
ing is applied to it. 

Taxonomic History 

Because the Sciurinae are active by day and often brilHantly 
colored, they have long claimed the interest of naturalists. In the 
Oriental Region, Horsfield (1824) was one of the earliest to attempt 
to summarize knowledge of the squirrels. In his "General enumera- 
tion of Indian Sciuri" he listed twelve species of non-flying squirrels 
under the generic name Sciurus, including representatives of the 
modern genera Callosciurus, Lariscus, Funambulus, and Ratufa. 

Gray (1867) briefly keyed out and roughly classified the Oriental 
squirrels by simple pelage and tooth characters but he missed what 
now seem to be the most obvious of relationships and lumped to- 
gether in one genus, Macroxus, some species now considered tribally 
different. John Anderson (1878, pp. 214-277) in order to identify 
his collections of squirrels from Burma reviewed all of the known 
Oriental squirrels. His conservatism in admitting Reithrosciurus 
and Rhinosciurus only to subgeneric rank seems a matter of the 
custom of his time, but the superficiality of his observations on the 
diagnostic characteristics of their skulls contrasts sharply with his 
penetrating observations on pelage color of squirrels in the Indo- 
chinese Subregion and his shrewd speculations as to the relationships 
that these indicate. 

Trouessart (1880) correctly recognized, as Gray (1867) had not, 
that the forms hicolor, giganteus, indicus, maximus, and macrurus 
belonged together in a taxon at least subgenerically distinct from 
typical Sciurus. He recognized Funambulus as of subgeneric value, 
but placed in it virtually every named Indochinese squirrel with 
stripes, including berdmorei, mcclellandi, and even quinquestriatus. 
In an Asiatic and Malaysian subgenus he also proposed to include 
erythraeus and lokroides with lokriah, pernyi, and rufigenus, among 
others, and included davidianus in the subgenus Rhinosciurus with 

Jentink (1883) recognized as belonging in the genus Sciurus the 
species erythraeus, lokroides, caniceps, atrodorsalis, quinquestriatus, 


castaneoventris, and pygerythrus from this region, but he included in 
the same genus the species lokriah and pernyi (now in Dremomys) 
and davidianus and berdmorei (now respectively in Sciurotamias and 
Menetes) and three species now properly belonging to Funamhulus. 
For the two subregions under discussion he admitted but one genus 
of diurnal squirrels and no subgenera. By grouping, however, he 
recognized affinities among species now known to belong to the genera 
Ratufa, Funamhulus, and Dremomys. 

Blanford's (1888-91) taxonomic treatment of the squirrels of 
these two subregions differs little from Jentink's, but Blanford pro- 
vided keys and more detailed characterizations. 

Major (1893, p. 189), using primarily characters of the teeth and 
to a lesser extent those of the skull, produced a classification of the 
living Sciuridae with six genera in the Sciurinae. One of the largest 
is Xerus, composed of five subgenera, four of which are entirely 
African tree and ground squirrels. The fifth subgenus was Eoxerus, 
and in it were the two commonest species of the Indian Subregion, 
palmarum and pennanti, now included in Funamhulus, herdmorei, 
now included in Menetes, and three Malaysian ground squirrel spe- 
cies of the present genus Lariscus. Under the genus Sciurus, sub- 
genus Sciurus, and beta group he included tree squirrels of the Indo- 
chinese Subregion — erythraeu^, atrodorsalis, caniceps, ferrugineus, and 
lokroides — and several tree squirrels of the Malaysian Subregion as 
well as one long-nosed species, everetti. Major ends this list with 
"etc.," which probably implies that long-nosed species rufigenu^, 
pernyi, and lokriah belong here, too. He holds to genus Sciurus, 
subgenus Eosciurus, for the several Oriental giant squirrel species 
as proposed by Trouessart. 

Thomas (1915) separated Indochinese and Malaysian tree squir- 
rel species from the genus Sciurus, noting that they are distinguished 
by the presence of a separate bony blade associated with the shaft of 
the OS penis. These tree squirrels of the Indochinese Subregion (and 
Malaysian Subregion) he recognized as Callosciurus and Tomeutes. 

Robinson and Kloss (1918) contributed a valuable list of the 
Sciuridae of the Oriental Region. They admitted nine species of 
Ratufa. In the Indochinese Subregion they admitted 15 species in 
what is now accepted as Callosciurus. 

Pocock (1923) followed Thomas' lead with a classification of the 
Sciuridae based upon further extensive study of the baculum (os 
penis) and glans penis. He associated the Indian striped squirrels, 
Funamhulus, and Oriental giant squirrels, Ratufa, with several genera 


of tree squirrels of the Ethiopian Region in one subfamily, but placed 
other Oriental squirrels (excluding Reithrosciurus) in a strictly Ori- 
ental subfamily, Callosciurinae. Simpson (1945) accepted Pocock's 
classification but scaled his subfamilies down to tribes. Moore (1959) 
found new skull characters which generally support Pocock's classi- 
fication as modified by Simpson, and by which one may allocate 
several Oriental genera, the bacula of which are still unknown, to 
the tribe Callosciurini. 

Ellerman (1940) offered an arrangement of the species and sub- 
species of Oriental squirrels, and Ellerman and Morrison-Scott (1951) 
offered an arrangement of species and subspecies of Sciurinae of the 
Indian and Indochinese subregions. These are checklists and not 
revisions at the species and subspecies level, and although there are 
many taxonomic decisions implicit in their arrangement, based on 
study of specimens, the evidence for these is not given. 

Zahn (1942) offered a revision of the genera, species, and sub- 
species of the giant squirrels, striped squirrels, and long-nosed squir- 
rels of the Oriental Region, based on study of material in European 

Other authors have published many helpful papers revising small 
groups of diurnal squirrels of this region. Some of these will be men- 
tioned in the following species accounts. Some authors have pro- 
vided important revisions of the Sciurinae included in large faunal 
papers. Of the latter the papers of G. M. Allen (1940) and Osgood 
(1934) are particularly helpful. 


The classification presented here constitutes conclusions derived 
from the whole present work and might with logic be placed at the 
end of the work. However, the choice had to be made between logic 
of presentation and convenience of use. We hope that some students 
will, indeed, weigh our logic, but we presume, however wrongly, that 
most students turning these pages, will be primarily interested in 
using our results. The attained classification is therefore provided 
in the introduction where it can serve conveniently as a framework 
for apprehending the work as a whole or the relationships of some 
particular genus or species. 

One of us (Moore, 1959) has reported in detail the tribal level 
skull characters that distinguish the following genera from all others 
in the Oriental Region: Ratufa (op. cit., p. 169), Funambulus (p. 170), 


and Sciurotamias (p. 182). The other existing genera of the Sciuri- 
nae of the Indochinese Subregion may be identified in the key to 
the genera and subgenera of the subtribe Callosciurina (op. cit., 
pp. 173-174). Tamiops is treated as a subgenus in that key. 

Classification of the Sciurinae of the Indian and 
Indochinese Subregions 

Tribe Ratufini Moore, 1959. Oriental Region. 
Genus Ratufa Gray, 1867. Oriental Region. 

Species macroura Pennant, 1769. Ceylon and tip of Indian Peninsula. 
Subspecies macroura Pennant, 1769 (including: 
ceylonicus Erxleben, 1777. 
ceilonensis Boddaert, 1785. 
tennenti Blyth, 1849. 
montanus Kelaart, 1852. 
macrurus Blanford, 1891.) 
melanochra Thomas and Wroughton, 1915. 
dandolena Thomas and Wroughton, 1915 (including 
sinhala Phillips, 1931.) 
Species indica Erxleben, 1777. Indian Subregion. 
Subspecies dealbata Blanford, 1897. 

indica Erxleben 1777 (including: 
purpureus Zimmerman, 1777. 
bombayus Boddaert, 1785. 
elphinstonei Sykes, 1831. 
superans Ryley, 1913.) 
maxima Schreber, 1784 (including: 
mxilabaricus Scopoli, 1786. 
bengalensis Blanford, 1897.) 
centralis Ryley, 1913. 
Species bicolor Sparrmann, 1778. Indochinese and Malaysian Subregions. 
Subspecies phaeopepla Miller, 1913 (including: 
celaenopepla Miller, 1913. 
marana Thomas and Wroughton, 1924.) 
leucogenys Kloss, 1916. 
sinus Kloss, 1916 (not mapped). 
smithi Robinson and Kloss, 1922. 
condorensis Kloss, 1920 (not mapped). 
felli Thomas and Wroughton, 1916. 
gigantea McClelland, 1839 (including: 
macruroides Hodgson, 1849. 
lutrina Thomas and Wroughton, 1916.) 
hainana J. A. Allen, 1906 (including 
stigmosa Thomas, 1923.) 

Tribe Funambulini Simpson, 1945. Ethiopian and Oriental Regions. 
Subtribe Funambulina Moore, 1959. Indian Subregion. 
Genus Funambulus Lesson, 1832. Indian Subregion. 
Subgenus Prasadsciurus, new subgenus 
Species pennanti Wroughton, 1905. Northern India, W. Pakistan, Nepal. 
Subspecies pennanti Wroughton, 1905. 
lutescens Wroughton, 1916. 
argentescens Wroughton, 1905. 


Species palmarum Linnaeus, 1766. Central and southern India and Ceylon. 
Subspecies palmarum Linnaeus, 1766 (includes: 
penicillatus Leach, 1814. 
indicus Lesson, 1835. 
gossei Wroughton and Davidson, 1919.) 
comorinus Wroughton, 1905. 
bellaricus Wroughton, 1916. 
robertsoni Wroughton, 1916. 
bengalensis Wroughton, 1916. 
favonicus Lindsay, 1926. 
kelaarti Blyth, 1851. 
brodiei Blyth, 1849. 

olympius Thomas and Wroughton, 1915. 
Species trisiriatus Waterhouse, 1837. Western Ghats. 
Subspecies tristriatus Waterhouse, 1837 (including: 
dussumieri Milne-Edwards, 1867. 
annandalei Robinson, 1917.) 
wroughtoni Ryley, 1913. 
numxirius Wroughton, 1916 (including 
thomasi Wroughton and Davidson, 1919.) 
Species layardi Blyth, 1849. Ceylon and southern India. 
Subspecies layardi Blyth, 1849. 

signatus Thomas, 1924, 
dravidianus Robinson, 1917. 
Species sublineatus Waterhouse, 1838. Ceylon and southern India. 
Subspecies sublineatus Waterhouse, 1838 (including: 
delesserti Gervais, 1841. 
trilineatus Blyth, 1849.) 
obscurus Pelzeln and Kohl, 1886 (including 
kathleenae Thomas and Wroughton, 1915.) 

Tribe Callosciurini Simpson, 1945. Indochinese and Malaysian Subregions. 
Subtribe Callosciurina Moore, 1959. Indochinese and Malaysian Subregions. 
Genus Callosciurus Gray, 1867. Indochinese and Malaysian Subregions. 
Species erythraeus Pallas, 1778. Indochinese Subregion west of Irrawaddy 
Subspecies erythraeus Pallas 1778 (including 
wellsi Wroughton, 1921.) 
erythrogaster Blyth, 1842 (including: 
punctatissimus Gray, 1867. 
nagarum Thomas and Wroughton, 1916. 
kinneari Thomas and Wroughton, 1916. 
crotalius Thomas and Wroughton, 1916.) 
bhutanensis Bonhote, 1901 (including 

crumpi Wroughton, 1916.) 
intermedius Anderson, 1879 (including 

aquilo Wroughton, 1921.) 
sladeni Anderson, 1871 (including: 
kemmisi Wroughton, 1908. 
rubex Thomas, 1914. 
midas Thomas, 1914. 
vernayi Carter, 1942.) 
bartoni Thomas, 1914 (including: 
shortridgei Thomas and Wroughton, 1916. 
fryanus Thomas and Wroughton, 1916. 
miliar di Thomas and Wroughton, 1916. 


caryi Thomas and Wroughton, 1916.) 
haringtoni Thomas, 1905 (including 
solutus Thomas, 1914.) 

Species flavimanus I. Geoff roy, 1831. Indochinese Subregion east of Irra- 
Subspecies quinquestriatus Anderson, 1871 (including: 

heehei J. A. Allen, 1911. 

imarius Thomas, 1926. 

Sylvester Thomas, 1926.) 
gordoni Anderson, 1871. 
shanicus Ryley, 1914 (including 

Igriseopedus Blyth, 1847.) 
hyperythrus Blyth, 1855. (not mapped, no localities) 
michianus Robinson and Wroughton, 1911 (including 

haemobaphes G. M. Allen, 1912.) 
zimmeensis Robinson and Wroughton, 1916 (including 

primus Allen and Coolidge, 1940.) 
thai Kloss, 1917. 
atrodorsalis Gray, 1842. 
siamensis Gray, 1860 (including 

tachin Kloss, 1916.) 
pranis Kloss, 1916. 
rubeculus Miller, 1903 (including 

youngi Robinson and Kloss, 1914. Unmapped. Extrater- 
hendeei Osgood, 1932. 
castaneoventris Gray, 1842 (including 

insularis J. A. Allen, 1906.) 
flavimanus Geoffroy St. Hillaire, 1831 (including: 

quantulus Thomas, 1927. 

contumax Thomas, 1927. 

dactylinus Thomas, 1927. 

pirata Thomas, 1929. 

bolovensis Osgood, 1932.) 
griseimxinus Milne-Edwards, 1867 (including: 

leucopus Gray, 1867. 

fumigaius Bonhote, 1907. 

vassali Bonhote, 1907. 

phanrangis Robinson and Kloss, 1922.) 
gloveri Thomas, 1921. 
bonhoiei Robinson and Wroughton, 1911. 
styani Thomas, 1894 (including: 

griseopedus Milne-Edwards, 1874. 

Herpestes [sic] leucuru^ Hilzheimer, 1905. 

Herpestes [sic] albifer Hilzheimer, 1906. 

canigenus Howell, 1927. 

woodi Harris, 1931.) 
ningpoensis Bonhote, 1901 (including 

tsingtanensis Hilzheimer, 1905.) 
thaiwanensis Bonhote, 1901 (including: 

centralis Bonhote, 1901. 

roberti Bonhote, 1901. 

nigridorsalis Kuroda, 1935.) 
Species ferrugineus F. Cuvier, 1829. Indochinese Subregion east of Irra- 
waddy (including keraudreni Lesson, 1830.) 


Species finlaysoni Horsfield, 1824. Indochinese Subregion east of Salween 

Subspecies finlaysoni Horsfield, 1824 (including 

partus Kloss, 1915.) 
folletti Kloss, 1915. 
trotteri Kloss, 1916. 
frandseni Kloss, 1916. 
albivexilli Kloss, 1916. 
harmandi Milne-Edwards, 1876 (including 

pierrei Robinson and Kloss, 1922.) 
germaini Milne-Edwards, 1867. 
nox Wroughton, 1908. 
cinnamomeus Temminck, 1853 (including: 

splendens Gray, 1861. 

herberti Robinson and Kloss, 1922.) 
annellatus Thomas, 1929. 
williamsoni Robinson and Kloss, 1922. 
menamicus Thomas, 1929 (1928). 
sinistralis Wroughton, 1908 (including: 

dextralis Wroughton, 1908. 

lylei Wroughton, 1908. 

grutei Gyldenstolpe, 1917.) 
boucourti Milne-Edwards, 1867 (including: 

leucogaster Milne-Edwards, 1867. 

leucocephalus Bonhote, 1901. 

floweri Bonhote, 1901. 

tachardi Robinson, 1916. 

prachin Kloss, 1920. 

rajasima Kloss, 1920. 

cockerelli Thomas, 1928.) 
boonsongi, new subspecies. 
Species caniceps Gray, 1842. Indochinese Subregion west of Mekong River. 
Subspecies concolor Blyth, 1855 (including: 

lancavensis Miller, 1903. 

erubescens Cabrera, 1917. 

telibius Thomas and Robinson, 1921 Extraterritorial.) 
adangensis Miller, 1903 (including: 

terutavensis Thomas and Wroughton, 1909. 

moheius Thomas and Robinson, 1921. 

mohillius Thomas and Robinson, 1921.) 
bimaculatus Temminck, 1853 (including: 

epomophorus Bonhote, 1901. 

davisoni Bonhote, 1901. 

sullivanus Miller, 1903. 

matthaeus Miller, 1903. 

lucas Miller, 1903. 

milleri Robinson and Wroughton, 1911. 

samuiensis Robinson and Kloss, 1914. 

nakanus Thomas and Robinson, 1921. 

mapravis Thomas and Robinson, 1921. 

panjius Thomas and Robinson, 1921. 

panjioli Thomas and Robinson, 1921. 

tacopius Thomas and Robinson, 1921. 

pipidonis Thomas and Robinson, 1921. 

iabaudius Thomas, 1922. 

hastilis Thomas, 1923.) 


casensis Miller, 1903 (unmapped). 

fallax Robinson and Kloss, 1914 (unmapped). 

domelicus Miller, 1903 (including 

bentincanus Miller, 1903.) 
caniceps Gray, 1842 (including: 

chrysonotus Blyth, 1847. 

inexpectatus Kloss, 1916. 

helgei Gyldenstolpe, 1917. 

helvus Shamel, 1930.) 

Species phayrei Blyth, 1855. Indochinese Subregion west of Salween (in- 
cluding: blanfordi Blyth, 1862, and heinrichi Tate, 1954.) 

Species inornatus Gray, 1867. Indochinese Subregion east of Mekong (in- 
cluding imitator Thomas, 1925.) 

Species pygerythrus Geofifroy St. Hillaire, 1832. Indochinese Subregion west 
of Irrawaddy. 
Subspecies pygerythrus Geoffroy St. Hillaire, 1832. 
janetia Thomas, 1914. 
owensi Thomas and Wroughton, 1916. 
Twearsi Bonhote, 1906 (including: 

bellona Thomas and Wroughton, 1916. 
virgo Thomas and Wroughton, 1916.) 
stevensi Thomas, 1908. 
blythi Tytler, 1854. 
lokroides Hodgson, 1836 (including: 
assamensis Gray, 1843. 
similis Gray, 1867.) 

Genus Tamiops Allen, 1901. Indochinese Subregion and Malaya. 

Species mcclellandi Horsfield, 1839. Indochinese Subregion and extraterri- 
torial subspecies in Malaya. 
Subspecies mcclellandi Horsfield, 1839 (including: 
pembertoni Blyth, 1842. 
manipurensis Bonhote, 1900.) 
collinus Moore, 1958. 
inconstans Thomas, 1920. 
kongensis Bonhote, 1901. 
barbei Blyth, 1847. 
leucotis Temminck, 1853 (including 
novemlineatus Miller, 1903.) 

Species rodolphei Milne-Edwards, 1867. Thailand, Laos, Cambodia. 
Subspecies rodolphei Milne-Edwards, 1867 (including: 
liantis Kloss, 1919. 
lylei Thomas, 1920. 
dolphoides Kloss, 1922. 
holti Ellerman, 1940.) 
elbeli Moore, 1958. 
Species svnnhoei Milne-Edwards, 1874. Yunnan, Szechwan, Burma. 
Subspecies smnhoei Milne-Edwards, 1874 (including: 
clarkei Thomas, 1920. 
forresti Thomas, 1920. 
spencei Thomas, 1921. 
russeolus Jacobi, 1923. 
olivaceiis Osgood, 1932.) 
vestittis Miller, 1915. 
Species maritimus Bonhote, 1900. Southern China, Vietnam. 


Subspecies maritimus Bonhote, 1900 (including: 
formosanus Bonhote, 1900. 
sauteri J. A. Allen, 1911.) 
monticolus Bonhote, 1900. 
hainanus J. A. Allen, 1906 (including: 
riudoni J. A. Allen, 1906. 
laotum Robinson and Kloss, 1922.) 
moi Robinson and Kloss, 1922. 
Genus Dremomys Heude, 1898. Indochinese and Malaysian Subregions. 
Species lokriah Hodgson, 1836. Nepal, Assam, northern Burma. 
Subspecies lokriah Hodgson, 1836 (including: 
bhotia Wroughton, 1916. 
subflaviventris Thomas, 1922.) 
garonum Thomas, 1922. 
pagus Moore, 1956. 

macmillani Thomas and Wroughton, 1916. 
Species pernyi Milne-Edwards, 1867. Assam to Chekiang above Tropic of 
Subspecies pernyi Milne-Edwards, 1867 (including: 
griselda Thomas, 1916. 
lichiensis Thomas, 1922. 
lentus A. B. Howell, 1927.) 
howelli Thomas, 1922 (including: 
mentosus Thomas, 1922. 
imus Thomas, 1922.) 
flavior G. M. Allen, 1912. 
senex G. M. Allen, 1912 (including 

modestus Thomas, 1916.) 
calidior Thomas, 1916 (including 

chintalis Thomas, 1916.) 
owstoni Thomas, 1908. 
Species rufigenus Blanford, 1878. Indochinese Subregion and Malaya. 
Subspecies rufigenis Blanford, 1878 (including: 
fuscus Bonhote, 1907. 
adamsoni Thomas, 1914. 
ornatus Thomas, 1914. 
laomache Thomas, 1921. 
belfieldi Bonhote, 1908. 
opimus Thomas and Wroughton, 1916. 
Species pyrrhomerus Thomas, 1895. Southern China, northern Vietnam. 
Subspecies pyrrhomerus Thomas, 1895. 

riudonensis J. A. Allen, 1906 (including 

melli Matschie, 1922.) 
gularis Osgood, 1932. 
Species everetti Thomas, 1890. Montane Borneo. (Extraterritorial) 
Genus Menetes Thomas, 1908. Indochinese Subregion. 
Species berdmorei Blyth, 1849. Indochinese Subregion. 
Subspecies berdmorei Blyth, 1849 (including 
amotus Miller, 1914.) 
peninsularis Kloss, 1919. 
moerescens Thomas, 1914. 
decoratus Thomas, 1914. 
consularis Thomas, 1914. 
mouhotei Gray, 1861 (including: 

pyrrhocephalus Milne-Edwards, 1867. 
koratensis Gyldenstolpe, 1917.) 


Tribe Tamiasciurini i Simpson, 1945. Nearctic Region and Oriental Region. 
Genus Sciurotamias Miller, 1901. Montane China, Sikang to Chekiang. 
Subgenus Sciurotamias Miller, 1901. Montane China, Sikang to Chekiang. 
Species davidianus Milne-Edwards, 1867. Montane Sikang to Chekiang, 
Subspecies davidianus Milne-Edwards, 1867 (including 
latro Heude, 1898.) 
consobrinus Milne-Edwards, 1868-74 (including: 
collaris Heude, 1898. 
saltitans Heude, 1898. 
owstoni J. A. Allen, 1909. 
thayeri G. M. Allen, 1913.) 
Subgenus Rupestes Thomas, 1922. Montane Yunnan and Sikang, China. 
Species forresti Thomas, 1922. Montane Yunnan and Sikang, China. 

1 While the present paper was in press. Black (1963) published his dissertation 
on the North American Tertiary Sciuridae (localities in the United States and 
Mexico; 12 genera, 5 with living species). From that study Black says he proposes 
classification of the 37 living genera of Sciurinae of the world. However, besides 
including names of some North American fossil genera. Black (1963, p. 123) 
proposed only two changes from the most recent previous classification (Moore, 
1959, p. 198-200). 

One change suggested raising the chipmunk taxon from subtribal to tribal 
level because ... "I feel that they differ from the marmots and the spermophiles 
as greatly as they do from the tree squirrels and are fully entitled to tribal status." 
No evidence, no discussion of relative merits, only opinion based on study of 
Nearctic fossils representing two of the eight tribes earlier known. The present 
writer studied all of the living genera of Sciurinae comparatively and quantita- 
tively, amassed a great deal of evidence, tabulated and discussed this, and con- 
cluded that in comparison with the marmots and with all other suprageneric 
taxa of living sciurines, chipmunks merit subtribal rank (Moore, 1959, pp. 162- 
166, 180-182). It is recommended that their proposed elevation to tribal rank 
be rejected. 

The other change offered by Black (1963) is dissolution of the generally 
accepted tribe Tamiasciurini. Black (1963, p, 125) reports that 3 of the 26 skulls 
of Tamiasciurus at the Museum of Comparative Zoology have only two transbullar 
septa per bulla, but he evidently did not read the method for counting these 
provided in the caption of Table 1 (Moore, 1959, p. 163). There were 3 speci- 
mens with only two pairs of septa in 138 specimens at the American Museum of 
Natural History (Moore, 1959, table 1). If we add these to the 6 that I now find 
in 361 specimens of Tamiasciurus in Chicago Natural History Museum, the total 
is 9 per 499, or less than two percent. For the record, totals of specimens in the 
latter two museums for 2, 2*^, 3 3^^, and 4 pairs of septa are respectively 9, 12, 
473, 3, and 1 individuals of Tamiasciurus. Black's (1963, p. 125) proposal to re- 
duce the rank of the Tamiasciurini seems to be based on too small a sample, 
faultily observed, for one character (number of transbullar septa), and on an un- 
justified complaint that the other character (high squamosal) "is extremely hard 
to evaluate." Consequently, I have retained Tamiasciurini. 

I placed Sciurotamias in the Tamiasciurini (Moore, 1959, pp. 182-184) only 
tentatively, trusting it will stimulate publication of pertinent new observations, 
and I am keeping it there in the present work. 

Genus RATUFA Gray 

Ratufa Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 273. 
Rukaia Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 275. 
Eosciurus Trouessart, 1880, Le Naturaliste, 2, no. 37, p. 291. 

Type species. — Ratufa, Sciurus indicus Erxleben;/?wA;am, Sciurus 
macrourus Pennant; and Eosciurus, Sciurus hicolor Sparrmann. 

Definition. — The genus Ratufa contains only the species ma- 
croura, indica, hicolor, and affinis and is endemic in the Oriental 

Diagnosis. — Consult Figure 1 for some of the following details. 
(1) The squamoso-alisphenoid suture lies about halfway between the 
third upper molar and the posterior margin of the base of the zygo- 
matic process of the squamosal. (2) The chambers of the auditory 
bullae are not crossed by transbullar septa. (3) The palatines are 
very short, so that the pterygoid fossa extends as far forward as the 
third molar and sometimes almost as far as the second molar. (4) The 
postorbital constriction is less than 0.80 of the interorbital breadth. 
(5) In fully adult individuals both the sagittal suture and the fronto- 
parietal suture ankylose completely, so that there remain no visible 
sutures on the dorsal surface of the skull from the anterior edge of 
the frontals to the posterior edge of the parietals. (6) In lateral 
aspect the lateral lip of the infraorbital foramen is vertical to the 
occlusal plane, or inchned with the top forward, and the top of it 
reaches the maxillo-premaxillary suture. 

The above six characters distinguish Ratufa (which constitutes 
the tribe Ratufini), from other Sciurinae of the Indian and Indo- 
chinese subregions as follows: Sciurotamias by 1, 2, 3, 4, 5, and 6; 
sub tribe Callosciurina by 1, 2, 4, and 5; and Funambulus by 1, 2, 3, 
4, and 5. 

Systematic History. — Gray's (1867, p. 273) original characteriza- 
tion of Ratufa is not meaningful today. It consisted of "Tail elon- 
gate, longer than the body and head; large-sized." The name Ratufa 
was evidently meant as a subgeneric designation, and included only 


Fig. 1. Skull and left mandible of the pied giant squirrel, Ratufa bicolor, 
AMNH No. 83440, X 1. Note generic characters described in text. For usage of 
skull anatomy consult Moore (1959, figs. 1-5). 



the species Sciurus indicus. Farther on in the same paper he placed 
giant squirrels macroura of Ceylon, bicolor of Java, and ephippium 
of Borneo in a subgenus Rukaia under a genus Macroxus, but ad- 
mitted no close relationship between indicus and the other three. 

When Robinson and Kloss (1918) published a list of the squirrels 
of the Oriental Region, they employed the nine polytypic species of 
Ratufa which were then currently accepted : macroura, indica, bicolor, 
notabilis, ephippium, affinis, gigantea, phaeopepla, and melanopepla. 
Hayman and Holt (in Ellerman, 1940) followed that arrangement 
22 years later. At the same time, however, Chasen (1940) reduced 
notabilis, ephippium, and melanopepla to subspecific status, and Zahn 
(1942) reduced phaeopepla, gigantea, and indica to subspecies. Eller- 
man and Morrison-Scott (1951) accept as species macroura on Cey- 
lon, indica in the Indian Subregion and bicolor in the Indochinese 
Subregion. None of these authors, except Zahn, gave reasons for 
their decision or characters distinguishing the species, and as will be 
shown, the arrangement we present with reasons for decisions and 
characters for species, differs from that of Zahn (1942). 


1. Crown sharply separated from nape by a distinct color band between ears. . .2 
Crown pelage color connected with that of nape 3 

2. Digits black and ear- tufts inconspicuous macroura 

Digits maroon or buff and ear-tufts conspicuous indica 

3. Light mark on thigh; no black mark on chin affinis [extraterritorial] 

No light mark on thigh; small black mark on chin bicolor 

Intrageneric relationships. — It is suggested by the key to the spe- 
cies above and by the species diagnoses which will follow, that ma- 
croura is more closely related to indica than to bicolor or ajffinis. 
This is what would be expected, of course, on geographical grounds, 
and we think that it could also be demonstrated in characters of the 
skull. It seems desirable to point it out, however, because of a gen- 
eral similarity of appearance between some forms of macroura on 
Ceylon and forms of bicolor on Java and Bali (see Moore, 1960, p. 13). 

We could find no discrete descriptive characters of the skull which 
distinguish geographically adjacent pairs of these species. Some 
trends are apparent, but variation in them is considerable. For in- 
stance, the zygomatic process of the squamosal possesses an elevated 
ridge which rises to somewhat of a peak, and in lateral view makes 
(in adults) a distinctive second superior process on the zygomatic 
arch. This characterizes the USNM macroura material, somewhat 
variably characterizes the AMNH indica material, occurs in nearby 



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Mt. Victoria specimens of R. bicolor gigantea but not in Mt. Popa 
ones a little farther away. In the rest of bicolor and in afinis this 
process does not appear to occur with frequency. 

This skull character might be taken as unexpected support for 
Zahn's (1942) recognizing macroura, indica, and gigantea as conspe- 
cific, but not hainana and stigmosa which he included (all grouped 
because of ear tufts). There is, moreover, in the ANSP material a 
Philadelphia Zoological Park specimen without data of origin, which 
is surely a cross between Ratufa indica and R. bicolor, for it displays 
some of the characters of each. However, since Hill (1939 and 1942) 
has easily bred both macroura and indica in captivity, it seems likely 
that this specimen results from a cross made in captivity and cannot 
be represented as a natural intergrade between indica and bicolor, 
suggesting that they should be treated as subspecies. In the diag- 
noses of the species bicolor and indica in the present paper we have 
offered a total of seven pelage characters which clearly distinguish 
these two as species. 

In the present study it has also become clear that the extraterri- 
torial Ratufa affinis is taxonomically the most distinct of the four 
species, although one can certainly say that indica is more spectacu- 
lar. The equatorial giant squirrel affinis is the only one of the four 
species with three pelage characters distinguishing it from all three 
other species. 

Rather little correspondence with the relationships of the species 
of Ratufa described above is seen in the exceedingly scanty knowledge 
of bacula of this genus. Dammerman (1931, p. 454) figured and de- 
scribed the baculum oi R. b. bicolor of Java as "a simple short bone of 
9.4-9.9 mm. length, slightly upcurved; the basal end expanded and 
hollowed; the apex narrowed and flattened horizontally." From the 
other end of the range of this species, upper Burma, Pocock (1923, 
fig. 20, E and F) figured a baculum of R. b. gigantea which is very 
similar to the one from Java, but has the distal, flattened, ventral 
surface provided with four rows of small bony spicules. Pocock 
(1923, fig. 20, G and H) illustrated a baculum of R. indica showing 
it to be strongly curved upward distally, spread at the curvature, 
tridentate at the tip, and grooved or hollowed on its anteroventral 
surface. Prasad's (1954, fig. 4, D, E, and F) excellent illustrations 
of a baculum of R. indica maxima from Coorg or Mysore agree with 
Pocock's indica (a zoo specimen with no locality data) in the char- 
acters of base, amount of curvature, and lateral spread at curvature. 
Prasad's indica, however, does not show a hollow in its anterior sur- 


face, ends in a single nob that is slightly bilobed and has rows of 
bony spicules. Hill (1936, fig. 6, E and F) illustrated the baculum 
of R. macroura [dandolena] of Ceylon, showing that it has the 90-de- 
gree bend, widening at the bend, and a tridenticulate tip. 

In sum it appears that although variable, the bacula of hicolor 
may be distinguished easily from the equally variable ones of in- 
dica. However, until more published examples reveal the extent of 
variation, the existence of a real distinction between the bacula of 
indica and macroura is much less certain. 

It must be added that Hill (1940) reports rather striking differ- 
ences between the glans penes of R. macroura dandolena and R. m. 
melanochra, and that his illustration of the glans of melanochra is 
disconcertingly more like that of Prasad's (1954, fig. 4, A, B, and C) 
R. indica maxima than that of R. macroura dandolena. Much needed 
are further scientific reports on the glans penes and bacula of these 
and other subspecies (with the capture locality of each specimen and 
the identifying characteristics of each captive precisely stated) to 
demonstrate whether genital characters support or deny the species 
difference between macroura and indica which we now consider well 
established by our present study of characters of the pelage. 

The body and skull dimensions presented in Table 2 are measure- 
ments made and recorded by one of us (Tate) entirely of type speci- 
mens. These are offered here not as anything faintly resembling 
adequate representation of the subspecies or species, but as a rough 
indication of the dimensions likely to be found in this genus within 
the Indian and Indochinese subregions. 

Ratufa macroura (Pennant) 

Definition. — Ratufa macroura is the Ceylonese giant squirrel, which 
shares the southern India range of R. indica maxima but occurs with- 
out a competing giant squirrel species in Ceylon. 

Diagnosis. — Ratufa macroura has the following pelage characters: 
(1) The crown is black or blackish. (2) The digits are black or black- 
ish. (3) The dorsal pelage of feet, ankles, and forearms is yellowish 
buff. (4) Ear tufts are present but small and short, extending no 
more than 5 mm. beyond the skin of the ear tips. (5) A light-colored 
band crosses between the ears, separating crown from nape. 

The above characters distinguish macroura from the other spe- 
cies of Ratufa as follows: indica by 2 (ankle part of), 3, and 4; hicolor 
by 3 and 5; affinis by 1, 2, 4, and 5. 



Fig. 2. Entire species range of the Sinhalese giant squirrel Ratufa macroura 
as revealed by material examined. Subspecies: A, macroura; B, melanochra; 
C, dandolena. 

Relationship to other species. — Zahn (1942) treated macroura and 
indica as one species. We know of no evidence of intergradation 
between macroura and indica, and Abdulali and Daniel (1952) in 
their study of geographic variation in the species indica evidently 
observed none. The occurrence of R. macroura in a considerable 
geographic area within the range of R. indica in southern India with- 
out evidence of intergradation thus far, requires that the evidence 
regarding status be considered in detail. 

There is in the locality records presented here for R. m. dandolena 
(fig. 2) and R. i. maxima (fig. 3) some fragmentary evidence that in 
southern India where populations of these forms are geographically 
sympatric, they may be ecologically segregated. The available ele- 
vations of Indian localities where R. macroura dandolena were col- 


lected are relatively low (1000, 2000, and 3000 feet) ; whereas those 
for R. indica maxima tend to be higher (3200, 3285, 3300, 3500, 3500, 
3500, 4000, 4200, 4500, 5000, 6100, and 6500 feet). Both species 
were taken in the Palni Hills, two examples of macroura at 3000 feet 
on the north slope, one of indica at 4000 feet on the north slope, and 
one of indica on the summit (at Kodaikanal which is 5500 feet). It 
appears therefore that the two species may be segregated by eleva- 
tion, a circumstance which is not surprising in pairs of closely related 
and geographically sympatric species, and which evidently minimizes 
interspecific competition, enabling the two closely related species to 
evolve further in the same geographic area. 

The light coloration of macroura makes it seem more suited to a 
habitat of dryer, perhaps deciduous forest, and the generally dark 
color and preponderance of black in R. indica suggest that it is better 
suited to living in darker, denser, evergreen rain forest high in the 
mountains or on the windward slopes of the Western Ghats. The 
geographic distribution of these two species on the southernmost 
part of the mainland is further suggestive of this sort of ecological 
segregation. Khajuria (1955) listed both R. i. indica and R. macroura 
dandolena as members of the "humid element" of the mammal fauna 
of the Deccan. In view of the evidence advanced here, it appears 
that R. m. dandolena may now be better regarded as belonging to 
the mammalian fauna's "semi-humid element" of Khajuria's classi- 

It may be noted that as revealed by Abdulali and Daniel (1952, 
color plate) the named forms of indica occupying the Western Ghats 
form something of a color cline in which the lightest form, which is 
most like R. m. dandolena, is the most distant from dandolena, and 
the darkest form, which is least like dandolena, is the one closest to 
and sympatric with dandolena. Less strikingly but correspondingly, 
the lightest-colored subspecies of the species macroura is the one 
which is sympatric with the very dark subspecies of indica, and the 
blackest subspecies of macroura is geographically the most distant 
from the area of overlap with the dark subspecies of indica. This 
status appears to represent the phenomenon known as character 
displacement described and discussed by Brown and Wilson (1956, 
p. 49). 

Intraspecific variation. — Varying sexual dimorphism in size is at- 
tributed by Phillips (1935) to the several subspecies of macroura. 
He finds the males larger than the females of the subspecies ma- 
croura in samples of three males and six females, and in foothills 


material of dandolena in samples of five males and four females. 
But in samples of five males and six females of subspecies melano- 
chra he found the females lai:ger, and in the largest samples (Phillips, 
1935, p. 224) of eleven lowland males and nine lowland females, that 
we here regard as dandolena, he found the sexes very similar in size. 
We suggest that the supposed sexual dimorphism is an artifact of the 
small samples, and that larger samples would show the sexes to be 
alike in size. 

The averages and maxima that PhilHps (1935, pp. 218-225) gives 
for body measurements of the several subspecies of macroura do sug- 
gest the existence of general size differences between the subspecies. 
The order of size is from melanochra, the largest, down through ma- 
croura to dandolena (see data in subspecies accounts of the present 
paper) . 

Ratufa macroura macroura (Pennant) 

Sciurus macrourus Pennant, 1769, Indian Zool., 1, pi. 1, and text. 
Sciurus ceylonicus Erxleben, 1777, Syst. Regn. Anim., p. 416. 
Sciurus ceilonensis Boddaert, 1785, Elench. Anim., 1, p. 117. 
Sciurus tennenti Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 600. 
Sciurus macrourus var. montanus Kelaart, 1852, Prod. Faun. Zeylon., p. 50. 
Sciurus macrurus Blanford, 1891, Fauna Brit. Ind. . . . Mammal., p. 374. 

Types. — Sciurus macrourus, not seen, from "Ceylon and Mala- 
bar," restricted by Phillips (1933) to "the highland jungles of the 
Central and Uva Provinces," Ceylon; ceylonicus, ceilonensis, mon- 
tanus, and macrurus are misspellings or alternative names proposed 
for earlier names given here; S. tennenti not seen, and not found by 
Robinson and Kloss (1918, p. 185). 

Material examined. — Pattipola, C. P. [Central Province], Ceylon, 
6210 feet (B.M.), two. 

Discussion. — This is said to be a highland form (Phillips, 1935). 
It is apparently known only from specimens from the one locality, 
and is distinguished in these specimens by white tips on the hairs of 
its black tail, and a grizzled gray line along the sides between the 
black back and yellowish venter. 

Dimensions. — Phillips (1935, p. 218) alleges sexual dimorphism 
in this form, and provides averages of measurements in millimeters 
of three males and six females: length of head and body, males 346, 
females 375, length of tail, males 371, females [384], length of hind 
foot, males 68, females 71; length of ear, males 27, females 25.3. 
He records weights of two females that were evidently each three 


pounds, four ounces. For the average length of tail for the six females 
above, Phillips (1935, p. 218) actually gives "365 mm. (15.1 in.)." 
This figure in millimeters is less than length of head and body in the 
females, which seems a little improbable, less than length of tail in 
the males, which seems a little improbable, also, and does not equal 
15.1 inches. The figure 15.1 inches does exceed Phillips' averages 
for length of body of the females and also length of tail in the males. 
Assuming 15.1 inches to be more correct, we have provided its equiv- 
alent in millimeters in brackets above. 

Ratufa macroura melanochra Thomas and Wroughton 

Ratufa macroura melanochra Thomas and Wroughton, 1915, Jour. Bombay 
Nat. Hist. Soc, 24, p. 36. 

Type. — BM No., adult female from Kottawa, elevation 
280 feet, Southern Province, Ceylon, collected April 12, 1913, by 
E. W. Mayor. 

Material examined.— "Fasdon, Corola," Ceylon (BM), one; "Ana- 
sigalla," Matugama, W. P. [Western Province] (BM), one; Kalutara, 
Matugama, Western Province (BM), one. 

Discussion. — This is apparently a geographic race occupying all 
the wet lowland area, which lies southwest of the highlands between 
the highlands and the sea (Phillips, 1935). It is like R. m. macroura 
in having a yellowish venter and a black back and tail, but differs 
from it by having no white-tipped tail hairs or gray line along the 

Dimensions. — Phillips (1935, p. 221) provides averages of meas- 
urements in millimeters of five males and six females: length of head 
and body, males 369, females 360; length of tail, males 387, females 
388; length of hind foot, males 74, females 73.5; length of ear, males 
25, females 27; weight, males 3 lb. 6 oz., females 3 lb. 2 oz. 

Ratufa macroura dandolena Thomas and Wroughton 

Ratufa macroura dandolena Thomas and Wroughton, 1915, Jour. Bombay 

Nat. Hist. Soc, 24, p. 36. 
Ratufa macroura sinhala Phillips, 1931, Ceylon Jour. Sci., Sec. B, 16, p. 215. 

Types. — Ratufa m. dandolena, BM No., adult female, 
from Wellawaya, Uva Province, Ceylon, 608 feet, collected July 6, 
1913 by E. W. Mayor, sinhala, BM No., adult male col- 
lected at Nikawewa, near Kantalai, North Central Province, 250 
feet, February 21, 1929 by W. W. A. Phillips. 


Material examined, from Ceylon. — "Mousa Kanda," Gammad- 
uwa, C.P. [Central Province] (BM), one; Pukpitiya, Gammaduwa, 
Central Prov., 2000 feet (BM), one; Nikaweratiya, N.W.P. [North- 
west Province] (USNM), three; Tinnpani [Tirappane], N.C.P. [North 
Central Province], 275 feet (BM), one; Bintenna, Eastern Province 
(BM), one, (USNM), one; Mankeni, Eastern Prov. (BM), two; 
Maha Oya, Eastern Prov. (BM), three; Wellawaya, Uva Prov., 608 
feet (BM), one; Kumbukkan, Uva (BM), one; Tellula, Uva Prov., 
300 feet (BM), one; Ranna, S.P. [Southern Province] (BM), one; 
Hambantota District, S.P. (BM), two; "Wawonia" (USNM), seven; 
"Candelay" (USNM), two. 

Material examined, from southern India. — Kombu, south Coim- 
batore (BM), one; Chettiri Range, Salem District, Eastern Ghats, 
2000 feet (BM), two; Kurumbapatty, Salem District, Eastern Ghats 
(BM), two; seven miles north of Gungwadoria, north slope Palni 
Hills, 3000 feet (BM), one; north slope of Palni Hills, 3000 feet 
(BM), two; Shrivilliputtur, Madura, southern India, 600-1000 feet 
(BM), four. 

Discussion. — This geographic race of Ratufa macroura occupies 
forests of the dry lowlands of Ceylon and also an area in southern 
peninsular India. It is distinguished from the black-backed, black- 
tailed other two subspecies by having dorsal pelage entirely brownish 
gray except for the black patch of the crown and varyingly a black- 
ish area across the shoulders "and middle line of rump." The tail 
hairs have longer light tips than do those of R. m. macroura and these 
produce a much grayer overlay that obscures the black part of the 
hairs. The short midventral pelage of tail is consistently whitish. 
Tails on which the pelage is old, worn, and curling at the tips may 
be bleached to the color of the venter (yellowish buff). This descrip- 
tion is based on Thomas and Wroughton (1915a, pp. 36-37) and notes 
on the United States National Museum material. 

Phillips (1935, pp. 221-223) regards dandolena as a subspecies 
confined to forests of the foothills of the central highlands of Central 
and Uva provinces, and distinct from sinhala which Phillips recog- 
nizes as occupying the dry lowlands of Ceylon. The characters that 
he attributes to dandolena on a sample of nine in his arrangement 
appear to us to describe intergrades between subspecies macroura 
and Phillips' subspecies sinhala. Since no character distinguishes 
Phillips' concept of dandolena from both the upland macroura and 
lowland sinhala, and the intergrade characteristics are not shown to 
be constant over a considerable geographic area, it seems better to 



recognize only the two subspecies, upland macroura and lowland 
dandolena including the foothills specimens as intergrades. Neither 
Zahn (1942, p. 13) nor Ellerman and Morrison-Scott (1951, p. 497) 
have followed Phillips' arrangement of dandolena and sinhala. 

Dimensions. — Phillips (1935, p. 224) provides averages of meas- 
urements in millimeters of 11 males and nine females from the low- 
lands of Ceylon: length of head and body, males 331.2, females 329.8; 
length of tail, males 330, females 350.6; length of hind foot, males 
64.2, females 65.2; length of ear, males 24.1, females 25.1. Weights 
of one male and one female, respectively, are 2 lb. 5 oz., and 2 lb. 8 oz. 




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Fig. 3. Species range of the Indian giant squirrel, Ratufa indica as revealed 
by material examined. Subspecies: A, dealbata; B, indica; C, maxima; D, centralis. 


Ratufa indica (Erxleben) 

Definition. — Ratufa indica is the principal giant squirrel species 
which occupies the mainland of the Indian Subregion. 

Diagnosis. — (1) The ears have maroon tufts which extend about 
20 mm. beyond the skin of ear tips, and which are not pointed but in- 
stead have a great many hairs about the same length. (2) There 
is a narrow, dark, usually maroon stripe extending ventroposteri- 
orly from the anterior edge of each ear, and separating the light- 
colored side of the face from the still lighter-colored bib on the side 
of the neck. (3) There is a contrastingly-colored line which passes 
in front of the ear and meets its fellow at the back of the crown be- 
tween the ears and separates crown from nape. (4) Some or all of 
the dorsal pelage of the sides, back, and tail is maroon (when the 
pelage is not badly worn and faded) ; the remainder is black. 

The above characters distinguish Ratufa indica from the other 
species as follows: macroura by 1, and 4; bicolor by 1, 2, 3, and 4; 
and affinis by 1, 2, 3, and 4. 

Discussion. — We are indebted to Abdulali and Daniel (1952) for 
their record and analysis of the relationships of pelage color differ- 
ences to geography in this species, and no deprecation of the value of 
their work is implied by our taking a stand for recognition of sub- 
species which are more strongly marked and have greater geographic 
range and fairly large areas of intergradation. See Figure 3. 

Moore (1960, p. 13) has drawn attention to how the distribution 
of this species bears on the Satpura hypothesis. 

Ratufa indica dealbata (Blanford) 

Sciurus indicua var. dealbatus Blanford, 1897, Jour. Bombay Nat. Hist. Soc, 
11, p. 299, plate A, fig. 1. 

Type.— B.M. No., adult male from Mahal, Bangs, In- 
dia, collected April 6, 1896, by R. C. Wroughton. 

Material examined, all from India. — "Khandeesh " Dangs (B.M.) 
one; "Mahal" Dangs (B.M.) one (topotype); Mheskatri Surat, 
Dangs (B.M.), one; Songadh, Nadsari District (B.M.), one. 

Type description. — Ratufa indica dealbata has the appearance of 
an albinistic indica. The dorsal coloration of type examined in 1951 
is white above, becoming yellowish white on the thighs. The ear 
tufts and anterior part of face are yellowish white. The tail is almost 
white except for a darker shade on the basal two to three inches, 
caused by dark subterminal bands on the hairs which occupy the 


greater part of the length of the tail hairs. The under parts are 
white with a trace of yellowish on the insides of the hind legs. 

Discussion. — The original recognition of the existence of this form 
came from Blanford's report on Wroughton's collection of three speci- 
mens from the Dangs and Wroughton's examination of a captive 
specimen from the same place. Although Blanford's description and 
color plate left no doubt as to the distinctiveness of these specimens, 
subsequent authors (e.g., Ellerman and Morrison-Scott, 1951) have 
understandably been reluctant to accept this material as character- 
istic of a population of giant squirrels inhabiting a particular geo- 
graphic region. Salim Ali however, definitely established the con- 
tinuing existence of a population of these very pale squirrels in the 
Dangs fifty years after their original discovery there by Wroughton 
in 1896. Salim Ali collected six specimens of them in two localities 
additional to the type locality of Mahal (Abdulali and Daniels, 1952). 
The area known thus far to be occupied by these cream buff giant 
squirrels remains small. Nevertheless, the fact that they occupy the 
northwest extension of the species range, and the striking character 
of their local color difference, seem to justify recognizing them as a 

Although the status of dealbata as a subspecies is accepted, some 
uncertainty remains about its color characters. The one principle 
character is well understood: the dorsal pelage is whitish or cream 
buff (or as Blanford says "pale rufescent sandy"). Blanford says 
that this becomes "slightly more rufous on the posterior portion of 
the body and on the outside of the hind limbs, and . . brown on the 
outside of the forelimbs and on the basal portion of the tail." (His 
artist confined this "brown" of the "forelimbs" to the hands, prob- 
ably correctly.) Whiter than the above were the "forehead, a band 
down the back, all the tail except the basal portion, and the lower 
parts. . . ." The tufted ears, Blanford said, were "bright rufus," 
and although that is how Blanford's colored plate showed them, 
when one of us (Tate) examined the type specimen in 1951, he re- 
corded the ear tufts as "yellowish white." Abdulali and Daniel 
(1952) who had four specimens, say "Ear tufts brown." 

Tate's notes indicate that the darkness of the basal portion of the 
tail is due to long, dark, subterminal bands on the tail hairs there. 

While conforming otherwise to the above description, the little 
colored figure of dealbata presented by Abdulali and Daniel (1952) 
shows the color patch between the ears as dark (brown) contrasting 


with the cream buff dorsal pelage. Abdulali writes (letter of March 6, 
1959), however, that "the colour plate is erroneous ... [in showing] 
a dark patch between the ears; . . . there is no white or pale patch 
between the ears [either], and the grizzled whitish of the nape and 
back is separated from the white of the forehead and snout by an 
almost straight line between the ears." 

Dimensions. — Blanford's (1897, p. 301) original description in- 
cludes measurements of a pair of adults of this subspecies taken in 
millimeters by R. C. Wroughton, who collected the animals: length 
of head and body, male 369, female 370; length of tail, male 417, 
female 408; length of hind foot, male 75, female 69. 

Ratufa indica indica (Erxleben) 

Sciurus indicus Erxleben, 1777, Syst. Regn. Anim., p. 420. 

Sciurus purpureus Zimmermann, 1777, Spec. Zool. Geogr. Quadr., p. 518. 

Sciurus bombayus Boddaert, 1785, Elenchus Anim., 1, p. 117. 

Sciurus elphinstoni Sykes, 1831, Proc. Zool. Soc. London, 1831, p. 103, 

Ratufa indica superans Ryley, 1913, Jour. Bombay Nat. Hist. Soc, 22, p. 436. 

Types and co-types — Sciurus indicus, lost; purpureus, lost; bom- 
bayus, lost; elphinstoni, BM Nos., two adult males 
marked "Dekkan, India," and, an old female from "West- 
ern Ghats, Dekkan, India," all collected by Mountstuart Elphin- 
stone; superans, BM No., an old female taken at Wote- 
kolli. South Coorg, December 28, 1912, by G. C. Shortridge. 

Material examined, all from western peninsular India. — Devikop, 
2000 feet. South Mahratta (BM), two; "Wotekolli," 2000 and 3200 
feet, southern Coorg (BM), three; "Makut," 250 feet, southern Coorg 
(BM), one; Ghatmatha, Satara District (BM), one; Helwak, Satara 
District (BM), one; Samasgi, 2000 feet, Kanara Border, southwest 
Dharwar (BM), two; Yellapur, 1800 feet. North Kanara (BM), one; 
Landa [Londa], Bombay Pres. (AMNH), one; "Kanara," southern 
India (BM), three; "India" (USNM), one. 

Pennant (1771, p. 281) described this squirrel well "from a stuffed 
skin in Doctor Hunter's cabinet" and said it "Inhabits Bombay," 
but he gave it no name. The scientific name indicus and the syno- 
nyms purpureus and bombayus cited above were all frankly proposed 
as scientific names for Pennant's Bombay squirrel, and all three of 
their Latin descriptions of it show some direct translation from Pen- 
nant's English description. The Erxleben description of indicus we 
translate, ". . . tail as long as body, with tip orange. Length of [head 
and] body 16 inches, tail 17. Tips of ears tufted. Head, back, sides, 


upper part of legs, thighs and tail dull purple. Inferior part of legs 
and thighs, with the belly, yellow . . ." 

Discussion. — Possibly from just south of the Bangs for something 
like 100 to 150 miles to Poona, but definitely at Bhimashankar, a 
point 40 miles directly east of Bombay, Abdulali and Daniel (1952) 
have demonstrated that the forest is occupied by giant squirrels that 
have hazel dorsal pelage instead of bay, and that have tails whitish 
distally for about half of their length. South-southwest of Bhima- 
shankar about 25 miles, those authors found at Khandala 11 out of 
12 specimens to be bay instead of hazel, but at Khandala an aver- 
age of 0.40 (0.30 to 0.52) of the tail was whitish. Farther south they 
found bay dorsal color continued and whitish tip to the tail averaged 
only 0.25. This evidence suggests intergradation between the cream 
buff dealbata and the bay indica, and Abdulali and Daniel (1952) 
included the hazel material in the subspecies indica. However, if 
the hazel color and half white tail should prove to be constant in 
further material from the Western Ghats from Bhimashankar up to 
the vicinity of the Dangs, there would be a case for recognizing the 
giant squirrel of this area as a subspecies, R. i. elphinstoni. Abdulali 
writes, however (letter of March 6, 1959), that the additional ma- 
terial received by the Bombay Natural History Society since his 
1952 paper was written had yet included "no hazel coloured speci- 
mens from between Bhimashankar and the Dangs . . ." 

Dimensions. — Riley (1913 p. 436) provides ranges of some meas- 
urements taken in millimeters on an unstated number of indica speci- 
ments from Dharwar and North Kanara: length of head and body, 
340-380; length of tail, 370-446; length of hind foot, 73-77; length 
of ear, 25-33; greatest length of skull, 68-74; basilar length, 53.5-59; 
length of toothrow, 14.3-15.5; length of diastema, 15.5-18; length of 
nasals, 24.5-26; and zygomatic breadth, 46-49.5. 

Southward in the range of R. i. indica, a sample of 14 specimens 
considered identical with R. i. indica in color, was described as a new 
subspecies (Ryley, 1913, p. 436), R. i. superans, because an unstated 
number of the sample was discretely and notably larger than an un- 
stated number of ordinary subspecies indica from Dharwar and N. 
Kanara. Abdulali and Daniel (1952) reported another sample of 
three large specimens from near Hassan, and accepted superans as 
a good subspecies. Ellerman (1940), Zahn (1942), and Ellerman and 
Morrison-Scott (1951) have also accepted this as a good subspecies, 
but we are doubtful. Size alone in some mammals can evidently 
change with density of population (Scheffer, 1955), and in absence 


of other differences "superans" is regarded here as atypical samples 
of the typical subspecies. 

Ratufa indica maxima (Schreber) 

Sciurus maximus Schreber, 1784, Saugethiere, 4, p. 784, pi. 217B. 
Sciurus malabariciis Scopoli, 1786, Del. Flora Fauna Insul., 2, p. 85. 
Sciurus indicus var. bengalensis Blanford, 1897, Jour. Bombay Nat. Hist. Soc, 
11, p. 303, plateB, fig. 2. 

Types. — S. maximus, MNHN No. 93, adult female, from coast 
of Malabar, India, collected by Sommerat; bengalensis, BM No., adult from "India." 

Material examined, all from southwestern peninsular India. — 
"Cotengady Estate," 3500 feet [Nelliampathi Hills, Palghat Distr.], 
Cochen (BM), four; Kodaikanal, top of Palni Hills, Madras (BM), 
one; "Anamaad," 3200 feet, Malabar (BM), one; "Kukkal, Shola," 
6100 feet (BM), one; near "Machur," 4500 feet (BM), one; "Otta- 
coolie Estate," 3300 feet, Malabar [Nelliampathi Hills, Palghat 
Distr.] (BM), one; Penmudi [Ponmudi], Travancore (BM), one; 
"Northern slopes Palni Hills," 4000 feet (BM), one; Trivandrum, 
Travancore (BM), one; "India" (MCZ), one; Kellengode [Kollan- 
god], 3500 feet, S. India (AMNH), two; Nelliampathi Hills, 3500 
feet (AMNH), one; Kil, Kotajiri, 5700 feet, Nilgiri Hills (BM), two; 
"Attikan," 5000 feet, Mysore (AMNH), one; "Mysore" (AMNH), 
four; Avalanche, 6500 feet, Nilgiri Hills (AMNH), one; Kalhatti 
[Falls], 4200 feet, Nilgiri Hills (AMNH), one; Mudumalai, 3285 feet, 
base of Nilgris (AMNH), two; Ootacamund, Madras, India (AMNH), 

Disctission. — Blanford's (1897) subspecies bengalensis was de- 
scribed from a single specimen with locality unknown. Ryley (1913, 
p. 436) reported a sample of 10 from Coorg to be identical in color 
with the type of bengalensis. This color seen in Blanford's color 
plate of the type, and the color plate of Abdulali and Daniel (1952) 
is different from indica only by having a black tail. Since bengalen- 
sis seems to be present in pure form only in a small area, and since 
the one character distinguishing it from indica is possessed also by 
the adjoining subspecies to the south, maxima, it is difficult to re- 
gard bengalensis as anything but a rather attenuated area of inter- 
gradation between subspecies indica and maxima. Obviously, it 
does not fit the characters of either of these, but it is intermediate. 

From fifty miles south of Coorg in the Nilgiri Hills the American 
Museum of Natural History has material from Ootacamund and 


Mudumalai which possesses the "bengalensis" color pattern, and 
Abdulali and Daniel (1952) report two other such specimens from 
the Wynaad Nilgiris. They report two further specimens from the 
Nilgiri Hills which differ from the hengalensis pattern only in having 
additionally a patch of black on the upper forelimbs (not the shoul- 
ders) but point out that this may perhaps be disregarded because it 
also occurs inconsistently in the subspecies indica. They then re- 
port that from Kotagiri, only ten miles east of Ootacamund, the 
material is colored in the centralis pattern (i.e. differing from typ- 
ical indica in having a black tail and separate black shoulder patches). 
From Avalanche, about 10 miles in the opposite direction from Oota- 
camund, and from Kalhatti [Falls], which lies between two localities 
for the hengalensis pattern, the American Museum has specimens 
which approach the maxima color pattern more closely, but the feet 
are bay, the black of the shoulders is discontinuous, and the Kal- 
hatti one has entirely red hind legs. (Incidentally, these approach 
Scopoli's type description of malabaricus, but could hardly be more 
obviously intermediate between all of these other bay-footed color 
patterns and maxima.) The point is made above that in the Nilgiri 
Hills the hengalensis color pattern intergrades through examples of 
the centralis pattern toward and very closely approaching the color 
pattern of maxima. Possibly they may mix so that all three pat- 
terns could be observed at one locality, or more than one pattern in 
one brood. If they do not, the details of intergradation would make 
an interesting field study. 

Pelage color. — From the Nelliampathi Hills less than 50 miles 
south of the Nilgiri Hills the American Museum of Natural History 
has three specimens, two from Kellangode, all of which possess the 
color pattern which is typical for Ratufa indica maxima. This pattern 
is fore feet, fore legs, and hind feet Light Ochraceous-Buff ; the shoul- 
ders and nape black entirely across from one upper fore leg to the 
other; tail, rump, and thighs entirely black with an incipient narrow 
black line extending forward in the middorsum; bib Light Ochra- 
ceous-Buff (XV) and face forward of the red subaural stripe about 
Apricot Buff (XIV); crown, subaural stripe, 20 mm. ear tufts, a 
small anterior portion of the nape, and the shanks of the hind legs 
Morocco Red; sides Ox-blood Red; ventral pelage (which is long and 
dense enough to hide the dark basal color of the hairs in two of the 
three) Warm Buff (XV); interaural patch Light Buff. 

Diagnosis. — The subspecific characters of maxima are: the con- 
tinuous black across the shoulders, the black rump, the light feet. 


and the black tail tip, in that order of importance. The specimen 
from the Palni Hills which Abdulali and Daniel (1953, p. 731) de- 
scribe as a variant of maxima does not therefore seem importantly 
different, Moore wrote Humayun Abdulali to ask if in Bombay 
Natural History Society material maxima always has a single, con- 
tinuous shoulder patch and centralis always two separate shoulder 
patches. He responded (letter of March 6, 1959) that this is so. 

Habits. — Hutton (1949, p. 691) recorded observations of maxima 
in the High Wavy Mountains about sixty miles southwest of Ma- 
dura, some of which contribute toward understanding of factors 
which may influence their distribution: "A very common animal 
throughout the [broad-leaved] evergreen forests. It is not found in 
the deciduous forest. . . . They often use the same nests for some 
years . . . keeping them repaired and adding to them each year. 
I have examined several nests. . . . Their average size was 2 feet in 
diameter and the inside looked as if it would hardly accommodate 
an adult squirrel, being barely a foot across. They bring forth two 
lots of young each year, . . . one lot in September and another lot in 
March, the average per litter being two. Once only have I seen 
three. The adults . . . have no sense of territory. . . . When the 
young are past the milk-diet stage, both parents help in the feeding. 
Quite often I have seen these squirrels foraging together; one on the 
ground collecting nuts, etc., which the one in the tree knocks down. 
Very occasionally one comes across a squirrel larder at the base of 
some tree, which may contain upwards of two measures of nuts and 
other hard fruit." 

Dimensions. — Wroughton (1910, p. 889) provides several meas- 
urements of an unstated number of R. i. maxima, perhaps only one: 
length of hind foot, 80; greatest length of skull, 77; basilar length, 
60; upper molar series, 15; length of diastema, 16.5; length of nasals, 
24; zygomatic breadth, 49; interorbital breadth, 30. 

Ratufa indica centralis Ryley 

Ratufa indica centralis Ryley, 1913, Jour. Bombay Nat. Hist. Soc, 22, p. 436. 

Type.— BM No., adult male from Bori, Hoshangabad 
Central Provinces, India, 1600 feet, collected February 13, 1912 by 
C. A. Crump. 

Material examined, all from India. — Amarakantak, source of Nar- 
budda River (BM), one; Bori, 1800 feet, Hoshangabad (BM), five 
(topotypes); Balapalli, 1000 feet, Palkonda Hills (BM), one; Kolle- 
gal, Billigirirangan Hills, Coimbatore (BM), two; Chamarajnagar, 


about 2500 feet, South Mysore (BM), one; "Gerung River, Manip- 
we" (BM), one; Koduru, 550 feet, Balapalli Range (BM), one; Luia, 
1000 feet, Chaibassa, Bihar and Orissa (BM), three; Sangajata, 
1300 feet, Chaibassa, Bihar and Orissa (BM), one; Dhain, 1400 feet, 
Hoshangabad (BM), two; no data (AMNH), two; Amraoti, Bastar 
[Berar] (AMNH), two. 

Pelage color. — The subspecies centralis as exemplified by the Am- 
raoti material is: Ox-blood Red on the dorsum, sides, ear tufts, and 
hind legs; black on the forelegs, shoulders (but separated by 30 mm. 
of red), and tail (which, however, pales at the tip to Light Buff XV) ; 
Cinnamon-Rufous (XIV) on the feet; Light Buff on the bib and in- 
teraural patch, but Buffy Brown (XL) on the side of the face; Liver 
Brown (XIV) on the crown. 

Dimensions. — Miss Ryley (1913, pp. 436-437) provides ranges in 
millimeters of body and skull measurements of an unstated number 
of her hypodigm of about 20 specimens of centralis: length of head 
and body, 309-343; length of tail, 382-433; length of hind foot, 72- 
79; length of ear, 25-30; greatest length of skull, 69.5-74; basilar 
length, 50-58; length of tooth row, 14-15; length of diastema, 14-16; 
length of nasals, 21.5-24; and zygomatic breadth, 42-46. 

Abdulali and Daniel (1952, p. 473) provide ranges and averages 
of two measurements in millimeters for 19 specimens of centralis: 
length of head and body, 300-380, average 340; length of tail, 390- 
450 (but an exceptional 270), average 405. 

Discussion. — This well-marked although variable subspecies oc- 
cupies low hills of central India, as its name suggests, and follows 
these eastward almost to Bengal. It also occurs in the Eastern 
Ghats, however, and southward along this chain to the state of 
Mysore and reaches the Western Ghats in the Nilgiri Hills where it 
intergrades in a confusing three-way mixture with maxima of the 
south and indica of the northwest. 

Habits. — At Sangajata and Luia, Bihar, collector C. A. Crump 
took 13 specimens and wrote the following comments (Wroughton, 
1915c, p. 107) : "This handsome squirrel is gregarious and one of the 
most locally distributed animals I have ever come across. At Luia 
it inhabits a piece of jungle perhaps a mile square, and outside of 
this it is useless to search for it. Near Sangajata the favorite haunts 
were patches of jungle near the river, and though I passed through 
much of the Forest here, nowhere else, outside of the spots men- 
tioned, did I see the large conspicuous nests made by this species. 
The call is a loud rattle. . . . This squirrel lives among the most 


lofty trees, it can take huge leaps, and is equally at home on a smooth 
trunk or scrambling among the slenderest twigs. I have never seen 
it lower than about 12 feet from the ground . . ." 

Ratufa bicolor (Sparrman) 

Definition. — Ratufa bicolor is the bibbed giant squirrel which 
ranges over the greater part of the Malaysian Subregion and Indo- 
chinese Subregion at least as shown in Figure 4. 

Diagnosis. — Ratufa bicolor has the following characters of the pel- 
age: (1) A large whitish or buff, bib-like area sharply marked off on 
the side of the neck is continuous with the color on the side of the 
face. (2) In most subspecies a broad, moustache-like, black line ex- 
tends from the black rostrum down and back through the vibrissal 
patch and the whitish or buffy pelage on the side of the face. (3) 
There is a small black spot in the whitish pelage of the chin, often 
divided midsagittally and finely into two parts. 

The above characters distinguish Ratufa bicolor from the other 
species of Ratufa as follows: affinis by 2 and 3; indica by 1, 2, and 3; 
macroura by 1, 2, and 3. 

Intraspecific variation. — In Table 2 some measurements of nine 
specimens of the species bicolor provide a bare indication of the 
dimensions one may find in species bicolor in the Indochinese Sub- 
region. The whole range of this species of giant squirrel, as revealed 
by the listings of Chasen (1940) and of Ellerman and Morrison-Scott 
(1951), extends from Nepal to Hainan, South Viet Nam, and Bali. 
No definition has previously been proposed for this broad concept 
of the species bicolor. It was somewhat of a practice in the past to 
include the forms south of Malacca Strait (bicolor, baliensis, palliata, 
etc.) in the species bicolor, but the mainland and insular forms north 
of that strait (peninsulae, phaeopepla, gigantea, etc.) were regarded 
as belonging in one or more separate species. Robinson and Kloss 
(1918, pp. 118, 195) thus recognized in what is now regarded the one 
species, several species separable at the Malacca Strait, the Isthmus 
of Kra, and along mainland lines less well defined. Although by 
1922 Robinson and Kloss had evidently come to consider the above- 
named forms conspecific, Ellerman still accepted their 1918 classifi- 
cation for his "Families and Genera of Living Rodents" in 1940. 
No one has advanced any justification for recognizing a species divi- 
sion at the Malacca Strait in the first place, nor for abandoning it in 
the second; and we have empirically sought evidence on this. 



Fig. 4. Distribution of the mainland subspecies of Ratufa bicolor which occur 
north of the Isthmus of Kra: M, phaeopepla; N, leucogenys; 0, smithi: P, felli; 
Q, gigantea; R, hainana. 

The pelage characters common to the forms of Ratufa bicolor 
southwest of the Malacca Strait are: (1) The dorsal body pelage is 
generally light tipped. (2) the tail hairs are generally light tipped. 
(3) The black moustachial line is obsolescent. Numbers 1 and 2 
provide striking contrast with the pelage characters of all the forms 
(of species bicolor) north of Malacca Strait except smithi. Pelage 
characters common to the forms under discussion on both sides of 
the Malacca Strait are: (1) The ventral body pelage is bicolored. 
(2) There is a contrastingly light bib on the face and neck. (3) A 


black triangular mark is found on the chin. (4) There is a light 
flash mark on the dorsal pelage of the fore legs. (5) A light mark 
also ornaments the hind foot. In view of the number and character 
of the differences and similarities, a better case is deemed to exist 
for regarding these forms as conspecific. 

North of the Malacca Strait another place where the pied giant 
squirrels were regarded as divided into geographically juxtaposed 
species is at the Isthmus of Kra, where melanopepla and phaeopepla 
seemed to come together (Miller, 1913, p. 25; Robinson and Kloss, 
1918, pp. 194-195; Ellerman, 1940, p. 389). The division of forms 
at the Isthmus of Kra was regarded as but doubtfully of species level 
by Robinson and Kloss (1918, p. 195), and restudy of the original 
series involved and consideration of new material leads us to con- 
clude that it is at best a subspecies division. See the discussion here 
in the account of subspecies R. b. phaeopepla. 

The next former alleged species division line farther north, which 
separated the alleged species phaeopepla and gigantea may be dis- 
tinguished in refined form on our map, Figure 4, separating subspe- 
cies gigantea and hainana from subspecies phaeopepla, leucogenys, 
and smithi. The differences that we find involved here are: (1) Ear 
tufts occur in gigantea and hainana but not in the other subspecies. 
(2) Flash marks occur on the fore legs of phaeopepla, leucogenys, and 
smithi, but not on the other subspecies. (3) The white of the bib 
and cheeks rises up in front of the eyes in gigantea and hainana but 
does not in the other subspecies. These taxonomic differences are 
discussed in further detail below, particularly in the subspecies ac- 
count of gigantea. One may note that much more taxonomic differ- 
ence has been found between these subspecies groups here than 
between those at the Isthmus of Kra. 

Relationships to other species. — Zahn (1942) has taken the present 
geographic line of separation as the one that parts the Indian species 
from the species bicolor. For our part, however, we find that separa- 
tion to be at the subregional boundary, which may loosely be said to 
be the barrier imposed by the Ganges River and its valley (but see 
Moore, 1960). The maroon species indica is separated from the pied 
species of giant squirrel across the subregional boundary by a more 
impressive assortment of characters, each of which has wide geo- 
graphic appHcabihty beyond the subspecies mentioned: (1) A promi- 
nent whitish or buffy mark separates the crown from the nape in 
indica centralis but not in bicolor gigantea. (2) A dark line descending 
from the ear divides the bib into cheek and neck components in cen- 


trails but not gigantea. (3) The ears of centralis have large Maroon 
tufts, but those of gigantea have small black tufts. (4) Light-colored 
pelage covers the feet, wrists, and ankles of centralis but in gigantea 
these are black. (5) Large areas of the fresh dorsal pelage are Maroon 
in centralis, but the back of gigantea is only black. (6) A black or 
blackish chin spot is present in gigantea but not in centralis. (7) A 
black or blackish moustachial line extends ventroposteriorly from 
the rostrum in gigantea but not in centralis. This line is not to be 
confused or equated with the facial stripe described above as char- 
acter number two, as Zahn (1942) did equate them. 

On the basis of the above differences we consider the range of the 
species Ratufa bicolor to go as far north and west as Nepal. We pre- 
sume that bicolor does not cross the Ganges River or intergrade any- 
where with Ratufa indica. We regard all of the giant squirrels of the 
Indochinese Subregion to belong properly to the species bicolor. The 
range of the species as indicated by the material we have examined 
extends from its northwestern locality in Nepal some 1500 miles 
eastward across Assam, upper Burma, southern Yunnan, Tonkin, 
and southern Kwangsi to Hainan, The northwest to southeast ex- 
tent of its range, including the extraterritorial part, is from Nepal 
to Bali. Indonesia, some 3000 miles. 

In the Academy of Natural Sciences collection at Philadelphia 
we found a giant squirrel specimen with characters which seem defi- 
nitely a mixture of those of species indica and bicolor. Nothing is 
known of its geographic origin, and it came to the Academy from the 
Philadelphia Zoological Garden. W. C. 0. Hill (1939, 1942) has re- 
ported successful breeding in both species macroura and species in- 
dica in captivity, and in view of the origin of the Academy specimen 
it seems likely that it represents a species cross between captive speci- 
mens of indica and bicolor. A less likely origin would be a relict 
patch of forest, from which no other museum specimens were avail- 
able to us, naturally inhabited by an interbreeding mixture of indica 
centralis and bicolor gigantea. 

Ratufa bicolor phaeopepla (Miller) 

Ratufa phaeopepla Miller, 1913, Smithsonian Misc. Coll., 61, no. 21, p. 25. 
Ratufa celaenopepla Miller, 1913, Smithsonian Misc. Coll., 61, no. 21, p. 26. 
Ratufa phaeopepla marana Thomas and Wroughton, 1924, Jour. Bombay Nat. 
Hist. See, 24, p. 227. 

Types. — Ratufa phaeopepla, USNM No. 124235, adult male from 
Sungei Balik, Tenasserim, Burma, collected February 25, 1904, by 


W. L. Abbott; celaenopepla, USNM No. 124149, adult male from 
Domel Island, Mergui Archipelago, Burma, collected January 26, 
1904, by W. L. Abbott; marana, BM No., adult female 
from Mt. Popa, dry zone of upper Burma, collected September 26, 
1913, by G. C. Shortridge. 

Material examined, from Thailand. — Me Ping River 195 miles N. 
of Bangkok (BM), one; Lai Yoke [Ban Sai Yoke] (BM), one; 20 
miles west of Kampengpet, 450 feet (BM), one; Me Yam Valley, 
N. Phre 185 meters (BM), one; Me Taw Forest, Raheng (BM), one; 
Bank of Me Ping River, south of Hkampenpet, 120 to 180 feet (BM), 
two; Me Wong River, 53 miles E. of Um Pang, 800 feet (BM), three, 
(AMNH), three; 28 miles E. of Um Pang, 1750 feet (AMNH), two; 
"Wang Pratart Farm," Kam Peng Pet Prov. (CNHM), three; Ban 
Tong Ting (USNM), one; Raheng (USNM), one; Borphloy, Kan- 
chanaburi (USNM), one; Klong Klung, Kamphaengphet (USNM), 
two, (CNHM), five; Doi Phu Kha (USNM), one; Pak Koh (NR), 
one; Koon Tan (NR), three. 

Material examined, from Tenasserim, Burma. — Bankachon, V.P. 
(BM), four; Mergui (BM), one; Thagyet, Little Tenasserim River 
(BM), one; Thowngyoh [Theng-gan-ngok] (BM), one; Maliwun 
(AMNH), one; Taok Plateau (AMNH), one; "Bankusun" (BM), 
two; Victoria Point (BM), one, (USNM), two; Kisseraing Island 
(BM), two, (USNM), one; King Island (BM), one; Domel Island 
(USNM), two; Sullivan's Island (BM), one; Red Point (USNM), 
two; Sungei Balik (USNM), six; Telok Besar (USNM), five. 

Material examined, from Upper Burma. — Kodugwe, Pegu Yoma 
(AMNH), one; Camp Pinmezali, Pegu Yoma (AMNH), two; Mt. 
Popa (AMNH), five, (BM), two; 20 miles N. of Toungoo (BM), one; 
30 miles N. to N.W. of Toungoo (BM), two; Kokkogon (AMNH), 
one; "Shenege Chaung, 1000 feet, Suiges Ruen," Mandalay (BM), 
one; Taho, Kareni (USNM), one. 

Dimensions. — From the describer's hypodigm of 16 specimens of 
phaeopepla, all collected by W. L. Abbott in southernmost Burma, 
we find the modal weight in 15 recorded on specimen tags by the 
collector to be four pounds. Only the pregnant female mentioned 
below weighed more, and the lowest weight, 3^ lb., is of an imma- 
ture male. The collector's measurements of phaeopepla were: length 
of head and body, 365-415 (average 386); length of tail, 390-500 
(466). Twelve of the 16 were males. The type of celaenopepla was 
a male weighing five pounds, according to W. L. Abbott's label. 
See weights reported for R. b. gigantea. 


Re-examination of Miller's phaeopepla series of 16 specimens used 
in the original description, reveals 11 that are mature on the basis 
of ankylosis and tooth wear. The 11 range from 73.2 to 77.6 and 
average 75.5 in greatest skull length. The peninsulae available to 
Miller were only three, admissibly mature by the same criteria, but 
brought up to 12 by three adults more recently obtained from Kuala 
Lumpur, one from Bangnara, and five from Ban Don, the series at 
the U. S. National Museum ranges from 68.3 to 74.0 and averages 
71.9. This difference from phaeopepla seems at least feebly to sus- 
tain Miller's proposition of a taxonomic break at the Isthmus of Kra, 
but we consider this to be at no better than the subspecies level. 

Pelage color. — The newer U. S. National Museum material from 
Bangnara and from Waterfall, Trong, Thailand, includes three speci- 
mens with molt lines across the back showing that the new pelage 
of peninsulae is much blacker than the freshest pelage of Abbott's 
phaeopepla series. This seems also to validate Miller's statement 
that fresh pelage color distinguishes phaeopepla from peninsulae; al- 
though the fresh pelage of a molting immature from Rumpin River 
in Miller's hypodigm of peninsulae seems to us no darker than some 
in his hypodigm of phaeopepla. 

Diagnosis. — Miller only distinguished celaenopepla from phaeo- 
pepla as having the fresh dorsal pelage much blacker. His type of 
celaenopepla is far darker than any U. S. National Museum phaeo- 
pepla, but none of the latter demonstrates by having a sharp molt 
line the ultimate blackness that may be reached by phaeopepla. 

Ratufa bicolor phaeopepla occupies the southern two-thirds of 
Burma, a range about 800 miles long. It must be noted that Miller's 
material was all from the very southern end of the range of phaeo- 
pepla, and since this range as we see it, borders upon that of the 
conspecific gigantea, felli, hainana, and leucogenys, it requires also to 
be carefully distinguished from those subspecies. Ratufa bicolor 
phaeopepla is distinguished from gigantea and hainana by possess- 
ing a buffy flash mark spread across the dorsal surface of its fore legs, 
and by lacking distinct ear tufts; from felli by the dorsal pelage be- 
ing blackish with no outstanding break in color from crown to tip 
of tail; from leucogenys by having bib and flash mark on fore leg 
more richly colored, Salmon Color (XIV) instead of the nearly white 
Cartridge Buff (XXX) ; from peninsulae by larger adult size. 

Discussion.— W. L. Abbott noted that the USNM No. 124247 
which he collected March 27, 1904, from Telok Besar in southern 
Tenasserim weighed 4 3/^ lb. and had but a single fetus in the uterus. 


It seems at least noteworthy that specimens from along the Me- 
wong River east of Um Pang [Ban Le Kathe], Thailand, have the 
pelage of their napes faded to form a prominent pale spot as is com- 
mon in leucogenys. Since the bibs and flash marks on the fore legs 
of three seem also somewhat light for phaeopepla, these may be con- 
sidered intergrades with leucogenys. The specimen from Borphloy 
[King Bo Ploei], Thailand, is likewise intermediate. 

Habits. — Field observations from the northern and southern lim- 
its of the subspecies range: "Local and not very plentiful on Mt. 
Popa, only occurring in the heavy evergreen forest near the top of 
the mountain." (G. C. Shortridge in Wroughton, 1915a, p. 472.) 
"Fairly plentiful, but more difficult to obtain in thick evergreen 
jungle than in the deciduous forests further north, as in other places 
these squirrels become rust coloured during the hot season. Weight 
3-4 lbs." (G. C. Shortridge in Wroughton, 1915b, p. 712.) 

Ratufa bicolor leucogenys (Kloss) 

Ratufa melanopepla leucogenys Kloss, 1916, Proc. Zool. Soc. London, 1, p. 43. 

Type. — BM No., old female from Lem Ngop, southeast 
Siam, collected January 15, 1915, by C. B. Kloss. 

Material examined, from southeastern Thailand. — Hinlap, 900 
feet (BM), one; Krabin CBM), one; Sakerat (NR), one; Chanta- 
boon (AMNH), one; Twenty miles west of Kempenpet (AMNH), 
one; Twenty -eight miles east of Um Pang (AMNH), two; Me Wong 
River, fifty-three miles east of Um Pang (AMNH), three; Klong Yai 
(USNM), two; "Huey Yang," Sriracha (USNM), one; Kao Sabab 
(USNM), four; Nong Dom Ta (USNM), one; Nongkhor (USNM), 
two; Nong Mong Muang (USNM), one; Hoopbon [Ban Hup Bon] 
(USNM), one; Bor Phloy, Kanjanaburi (USNM), one; Ban Nong 
Bua (CNHM), one; Lem Ngop (CNHM), one. 

Material examined, from Cambodia. — Mt. Pra (BM), one; "Diai, 
Melton" (BM), one; Samba [Sambor] (BM), one. 

Original description. — "Characters. — Like R. m. peninsulae . . . 
but yellow of cheeks, fore limbs, and under surface markedly paler 
than the respective areas in that form, yellow on thighs more exten- 
sive and continued along the sides of the feet onto their upper sur- 
faces, where it occupies a considerable area, while the yellow of the 
fore limb extends to the bases of the toes above." 

Pelage color. — We made the following color notes from a sample 
of six specimens in the United States National Museum, and the 


Ridgway color terms used by Kloss are added in parentheses: bib 
and flash mark on fore leg are Cartridge Buff (Ivory Yellow) ; venter 
Apricot Buff (Pale Orange-Yellow). 

Diagnosis. — R. b. leucogenys differs from phaeopepla in having a 
paler bib and flash mark on the front leg, and in having a larger pale 
mark on the hind foot; from hainana by possessing a flash mark on 
the front leg and lacking ear tufts; from smithi by having no whitish 
tips on the dorsal pelage. 

Dimensions. — Measurements of three adults provided by Kloss 
in millimeters in the original description: length of head and body, 
370, 360, 345; length of tail, 435, 435, 450; length of hind foot, 79, 
80, 79; greatest length of skull, 71, 73, 71; condylobasal length, 60.5, 
61, 58.5; palatal length, 27, 27.7, 26.3; diastema, 16, 16, 14.7; length 
of maxillary toothrow, 14, 14.2, 14; interorbital breadth, 26.5, 29.5, 
26.7; zygomatic breadth, 46, 44, 44.5. Sex is respectively female, 
male, male. 

Ratufa bicolor sinus (Kloss) 

Ratufa melanopepla sinus Kloss, 1916, Proc. Zool. Soc. London, 1, p. 44. 

Type.— BM No., adult female from Koh Kut Island, 
southeast Siam, collected December 26, 1914, by C. Boden Kloss. 

Material examined. — Koh Kut Island, southeast Siam (BM), one, 
(USNM), three. 

Diagnosis. — This subspecies is most like leucogenys to which it is 
geographically adjacent, and differs from it by having uniformly 
clear black dorsal pelage and richer colored ventral pelage, about 
Orange-Cinnamon we note, but Kloss describes it as varying from 
Ochraceous Buff to Ochraceous Orange and Ochraceous Tawny in 
the center of the abdomen. He noted that the nasal bones were 
rather longer in sinus than in leucogenys, their posterior terminations 
being more in line with those of the premaxillaries. 

Dimensions. — The original description provided measurements in 
millimeters of six adults, three of each sex. Ranges and averages of 
these are: length of head and body, 355-380 (366); length of tail, 
415-450 (431); length of hind foot, 75-79 (77); greatest length of 
skull, 71-73 (71.9); condylobasal length, 59.5-61.2 (60.3); palatal 
length, 27-28 (27.3); length of diastema, 15.5-16.3 (15.8); length of 
maxillary toothrow, 14-14.2 (14.1) ; interorbital breadth, 28-30 (28.5) ; 
zygomatic breadth, (five specimens) 44^6 (45.1). 

Ratufa bicolor sinus is not mapped in Figure 4. 


Ratufa bicolor smithi Robinson and Kloss 

Ratufa bicolor smithi Robinson and Kloss, 1922, Ann. Mag. Nat. Hist., [ser.] 9, 
9, p. 89. 

Type.— BM No., old female from Langbian Peaks, 
south Annam, 6000 feet, collected April 22, 1918, by C. Boden Kloss. 

Material examined. — Tay Ninh, Cochin China (BM), one; Tay- 
ninh Mt., Cochin China, 1000 meters (MNHN), one; Djiring, south 
Annam, 3500 feet (BM), three; Ban Me Thuot, Annam (CNHM), 
two; Langbian Peaks, south Annam (USNM), one. 

Discussion. — This is a strikingly distinct subspecies with a small, 
poorly known range in southern Vietnam. The describers had ex- 
amined nine specimens from three localities: Arbre Broy^ at 5400 
feet elevation, Dalat at 4500 feet, and Dran at 3000 feet in the Lang- 
bian Mountains. It is notable that none of these localities, or of 
those from which we have examined material, is from below 3000 
feet elevation, and that the greatest diameter of its known range 
barely exceeds 150 miles. 

Pelage color. — The characters which distinguish it are: buffy tips, 
to as long as 15 mm., on the hairs of the back, producing about Light 
Ochraceous Buff in USNM No. 269797; the light tipping of the hairs 
intensifies to produce a distinct patch on the posterior part of the 
crown, about Light Buff in the same specimen; the pelage of the 
sides, limbs, and tail is black when the pelage is fresh. The buffy 
tipping extends onto the thighs and base of the tail to a small extent 
and sometimes onto the forelimbs. The black parts do fade to brown 
to some extent, particularly toward the tip of the tail. The color of 
the hair tips on the back varies, probably from fading and wear, but 
the molt line across the back of USNM 269797 shows very little 
color change from old pelage to new. Ventral pelage Antimony Yel- 
low to Warm Buff, according to the describers, and bib, flash mark 
on fore legs, and frequently the hind feet, are the same color. 

Diagnosis. — Closely related to leucogenys, particularly by the pale 
nape patch, but also by the flash mark on the fore leg and the hind 
foot color, smithi is nevertheless sharply distinguished from it by the 
light-tipped pelage running the length of its back. From hainana, 
smithi differs by possession of the flash mark on the fore legs and by 
lacking ear tufts. 

Dimensions. — Kloss' measurements in millimeters of the type are: 
length of head and body, 415; length of tail, 500; length of hind foot, 
89; length of ear, 31. The describers say that the skull of the largest 


of nine specimens measured 79 and 50 mm. in greatest length and 
breadth of skull respectively. 

Ratufa bicolor condorensis (Kloss) 

Ratufa melanopepla condorensis Kloss, 1920, Jour. Nat. Hist. Soc. Siam, 4, 
p. 71. (Not R. condurensis Miller, 1907.) 

Type. — C. Boden Kloss No. 2706 (not found by the present re- 
visers), adult female from Main Island, Pulo Condore, off the coast 
of South Viet Nam (not Pulo Condur of the Malacca Straits). 

This insular form shows close relationships to leucogenys and 
smithi in consistent possession of a light nape patch. It differs from 
both of these primarily in size, for condorensis is a pigmy giant 
squirrel, the skulls of seven specimens measuring 60, 61.5, 62, 62.5, 
62.5, 63, 64mm. A greatest skull length of smithi (Robinson and Kloss, 
1922, p. 93) is 76 mm., and for three leucogenys is 71, 71, 73 mm. 
(Kloss, 1916, p. 69). This small insular form differs from smithi by 
lacking light tips to the pelage of the back, but otherwise resembles 
both smithi and leucogenys closely in pelage color. 

No material representing this subspecies was examined during the 
present study. This subspecies is not shown on the map in Figure 4. 

Ratufa bicolor felli (Thomas and Wroughton) 

Ratufa felli Thomas and Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, 
p. 226. 

Type. — BM No., adult male from Yin, East bank of 
Lower Chindwin, Burma, collected June 13, 1914 by G. C. Short- 

Material examined, all from Burma. — Dudaw-Taung, 650 meters, 
Pakokku Chin Hills (AMNH), two; Ainggyi, Pakokku Chin Hills 
(AMNH), one; Mt. Victoria, 500 meters (AMNH), one; Yin, Lower 
Chindwin (BM), 4; Okma, east bank of Chindwin R., N. Burma 
(AMNH), two. 

Pelage color. — This appears to be a well-marked subspecies with 
a small, and poorly known, geographic range. It differs from gigantea 
and hainana to the north by having no ear tufts, by having an inva- 
sion of buffy from the insides of the fore legs virtually across the dor- 
sal surface, which is otherwise black, by having a more pronounced 
border of the ventral pelage. It shares with hainana a somewhat 
richer ventral pelage color than that of gigantea, and also the light 
mark on the side of the hind foot, and it shares with peninsulae to 


the south, all of these characters. It differs from all of its conspecific 
geographic neighbors, however, in that its dorsal pelage from the 
shoulders to the rump and the pelage of its sides from the fore legs to 
the hind legs is pale even when fresh. This ^vesfelli a general dorsal 
color pattern like that of Ratufa bicolor palliata of Sumatra: black 
on the head, nape, limbs, and tail, and light brown between the fore 
and hind quarters. In the present subspecies, however, the color of 
the sides is just like that of the back, not distinctly lighter as is the 
case in palliata. In one example of felli with apparently fresh glossy 
pelage the middle dorsal area is very close to but a little browner than 
Light Ochraceous Buff (XV), whereas in another it is about Cinna- 
mon-Buff (XXIX). The head and limbs are about Blackish Brown 
(1) XLV, and the nape and rump are intermediate. The intermedi- 
ate pale brown of the rump continues out the tail for about a deci- 
meter in the two with fresh pelage. The ventral pelage is blackish 
at the bases of the hairs on the hind legs as well as on the body, and 
is tipped with Cream Color to Warm Buff on an individual. The 
bib and blaze on the fore leg is Cream Color. 

It must be noted that the Mt. Victoria specimen is black dorsally 
like gigantea, but has no ear tufts, and is intermediate in the extent 
of the white blaze or flash mark across the fore leg, in having a small 
light mark on the side of the hind foot, and in the intensity of color 
of the ventral pelage. This specimen is either an intergrade between 
felli and gigantea or a challenge to the validity of felli as a subspecies. 
In view of the striking character of the felli material, we assume the 

Ratufa bicolor gigantea (McClelland) 

Sciurus giganteus McClelland, 1839, Proc. Zool. Soc. London, 1839, p. 150. 
Sciurus macruroides Hodgson, 1849, Jour. Asiatic Soc. Bengal, 18, p. 775. 
Ratufa gigantea lutrina Thomas and Wroughton, 1916, Jour. Bombay Nat. 
Hist. Soc, 24, p. 226. 

Types. — Sciurus giganteus, BM No., young adult 
from Upper Assam, India, collected by J. McClelland; macruroides, 
BM No., adult cotypes from Nepal collected in 1830 
and 1840 by Hodgson; lutrina, BM No., old female from 
Tatkon, near Kindat, Upper Chindwin River, Burma, collected 
July 5, 1914 by G. C. Shortridge. 

Material examined, from easternmost India. — Duragiri, 1600 feet, 
Garo Hills (BM), one; "Langting," 1500 feet, Cachar Hills (BM), 
two; Mokokchung, 4500 feet, Naga Hills (BM), one; "Naga Hills" 


Assam (BM), three; "Lakhani," Naga Hills, Assam, 1000 feet (BM), 
one, (AMNH), one; Tura, Assam (AMNH), two, (CNHM), two; 
"Chang chang Pani," Assam, 900 feet (AMNH), two; Ukhrul, Mani- 
pur, 6000 feet (BM), 1; Nongpoh, Khasi Hills, Assam (AMNH), 1, 
(CNHM), one; "Nepal" (BM), three; Pemionchee, Sikkim, 7000 feet 
(BM), one; Matanga River, 2500 feet, N. Kamrup (BM), two; 
Pashok, 3500 feet, Darjeeling (BM), one; Chin Hills 35 miles west 
of Kindat, 1500 feet (BM), one; Sevoke, Bengal (CNHM), two; 
Sangsir, 1400-2000 feet, Bengal (CNHM), three, (USNM), two; 
Tarkhola, Sikkim (CNHM), two; Karong, Manipur (CNHM), one. 

Material examined, from Tibet. — "Dening," 2250 feet, Mishmi 
Hills (BM), one; "Piki/' Mishmi Hills 1440 feet (BM), one; 
"Drexi," Mishmi Hills, 5140 feet (BM), one. 

Material examined, from Burma. — "Haingyan," 5000 feet. Chin 
Hills (BM), one; Hkamti, 500 feet, west bank upper Chindwin (BM), 
two; Kachin Hills 100 miles north of Myitkiyna, 1500 feet (BM), one; 
Moungkan, between Homalin and Tamanthe, east bank upper Chind- 
win River, 420 feet (BM), one; Lonkin (AMNH), one; Heinsun, 
west bank Chindwin R. (AMNH), one; Linhpah, west bank Chind- 
win R. (AMNH), one; Kindat (BM), one, (AMNH), one; Kabaw 
Valley, 20 miles west of Kindat, 600 feet (BM), one; Taukchaun, 
east bank of Chindwin, 500 feet (BM), one; Tatkon (BM), two; 
Road from Tamanthe to Sinnaing, west bank Chindwin R. (AMNH), 
one; Moungkan east bank Chindwin R. (AMNH), six; Hualung, 
east bank Chindwin R. (AMNH), two; 25 miles W. of Myitkyina 
(USNM), three; "Sedaw, Mandalay Canal and District," 300 feet 
(BM), one; Kamaing, Myitkyina Dist. (BM), one; N'bunghku 
(AMNH), one; Haibum (AMNH), four; Pumsin (AMNH), one; 
Htingwan, 4000 feet (BM), two; Hkani, 3000 feet, Gaw (BM), one; 
Singkaling Hkamti (AMNH), three. 

Pelage color. — From Nepal eastward across Sikkim, Assam, north- 
ern Burma, Yunnan, northern Thailand, and northern Indochina 
to Hainan the giant squirrel is characterized by tufted ears and en- 
tirely black dorsal pelage on the fore legs. In Assam and northern 
Burma we find that considerable American Museum of Natural His- 
tory material shows the tips of the ventral body pelage to be rather 
consistently pale, ranging between Cartridge Buff (XXX) and Light 
Buff (XV) on an individual (judged in places where the dark gray 
of the hair bases is entirely hidden by the pale tips). At Maymyo, 
Wan Tien, and the Nam Ting River, however, the ventral pelage 
shows enrichment at least to Light Ochraceous Buff, and this is true 


of the material from northern Thailand. This more intense color is 
here considered to be but a dilution of the still richer color charac- 
terizing the ventral pelage of the subspecies described from Hainan, 
which is consistently Apricot Buff to Ochraceous Orange on each 
Hainan individual. 

Thomas (1923, pp. 85-86) has shown that in northern Thailand 
the giant squirrel has a little patch on the side of each (otherwise 
quite black) hind foot that is the color of the ventral pelage. He 
found that this character diminishes greatly in northern Burma and 
Assam, and is entirely absent in the Mishmi Hills, Sikkim, and 
Nepal. We note, however, that it continues strong from northern 
Thailand to the westward and characterizes the subspecies which 
had been already described from Hainan, and so is not diagnostic 
for the subspecies stigmosa which Thomas (loc. cit.) erected upon it 
in Thailand. We consider that this character like that of the rich 
color of the ventral pelage, may emanate from Ratufa bicolor hainana. 

Diagnosis.— Consequently, Ratufa bicolor gigantea may be iden- 
tified by presence of ear tufts, black dorsal pelage on the forelegs, 
pale ventral pelage (about Cartridge Buff to Light Buff), and scarcity 
of any buff pelage on the side of the hind foot. The Maymyo and 
Nam Ting River specimens are apparently intergrades with hainana 
to the southeast. 

Variation. — Possibly there is a geographic race which should be 
recognized as Ratufa bicolor lutrina, occupying an area in northern 
Burma. However, the characters on which lutrina has been based 
are virtually identical with the faded condition of the pelage of some 
undoubted gigantea just prior to molt. If the brown color of lutrina 
were the color of the fresh pelage, and if it predominated in a geo- 
graphic area, we would freely accord subspecies status to lutrina as 
have Zahn (1942), Carter (1943), and Ellerman and Morrison-Scott 
(1951). There could, of course, quite by chance be taken in a given 
area. A preponderance of specimens whose pelage had come in black 
when fresh but had faded to brown at the time of collection. There 
could also be a greater tendency in a geographic area for the pelage to 
fade quickly to brown, so that it is brown for a longer time before 
being shed and replaced by black. This quick change to brown could 
be effected by a special genetic factor or a special ecological factor or 
both in combination. Carter (1943) considered that this frequency 
of brown pelage was related to the animal's occurrence in the savan- 
nah forest, and he allocated the blacker specimens that he studied 
from the adjacent more heavily forested districts to the subspecies 


gigantea. This necessitated arbitrary placing of a series from the 
deep forests of Lonkin and Singkaling Hkamti in gigantea even 
though the series is almost as brown as the alleged lutrina. Since 
there are small spots of new black pelage coming in in two of the 
brown specimens from Maunkan, the general brown color of the dor- 
sal pelage of these "lutrina'' is evidently produced by fading, and on 
the basis of the scanty information on its distribution now at hand, 
we give no credence to "lutrina" as a valid subspecies. 

There is, further, some individual variation in pelage color with- 
in gigantea which should be mentioned. Near the base of the tail 
for about a quarter or a third of the tail length the short, appressed 
ventral hairs of the tail are usually buffy instead of black. In one 
specimen from Tura this buffy line extends for two-thirds the length 
of the tail. This specimen also has reddish subterminal color bands 
on the hairs of the dorsum and tail which contrast strongly enough 
with the basic blackish brown to be quite noticeable. Further, it 
has a splash of white-tipped hairs in the black dorsal pelage of each 
fore leg (about 15 mm. in diameter) and a whitish tipping to the hairs 
on one side of the hind foot. Another specimen from Tura has none 
of these variations from proper gigantea characteristics. A specimen 
from Nongpoh in the Khasi Hills is quite as brown as any of the 
alleged lutrina, and has even the pale tail tip. A specimen from 
"Changchang Pani," Assam, is richly colored enough ventrally and 
possessed of a strongly enough marked hind foot to be an intergrade 
with hainana were that geographically possible. One Hualung speci- 
men from the east bank of the Chindwin River has almost whitish 
subterminal bands on many of the hairs of the dorsum, creating a 
rather agouti effect. 

Habits. — In Sikkim collector C. A. Crump took 14 specimens at 
six locaHties and wrote these notes (Wroughton, 1916a, p. 486): 
"This squirrel is usually found in pairs and is not gregarious. It is 
more plentiful between the Terai and the low valleys up to 3,000 feet, 
being particularly partial to the dense tall jungle bordering rivers. 
It has a loud clacking note common to most squirrels but calls only 
on rare occasions. It is an extremely active chmber and as a rule 
is wary, making off and hiding in the thick foliage immediately dan- 
ger is feared." 

Dimensions. — Collector H. Stevens noted weights of the three 
adult squirrels from Sangsir and Sevoke, Bengal Residency, India, 
on the specimen tags as four, four, and five pounds. These are all 
males taken in November and December, and are CNHM Nos. 


35432, 35434, and 35437. This may be the giant squirrel's best 
claim to being the largest tree squirrel in the world. He found the 
female specimen from Tarkhola in the Teesta Valley of Sikkim to 
weigh 43^ pounds. 

Discussion. — In January, 1931, Lord Cranbrook journeyed north- 
west from Putao, Burma, across the drainage of the Mali Hka to the 
Nmai Hka. While in the drainage of the former he heard (letter to 
us of May 5, 1961, based on his field notes) or saw giant squirrels 
daily. But once over the 6500 foot divide into the Nmai Hka Val- 
ley, he saw no more Ratufa; although "the vegetation and climate 
seem to be exactly the same on both sides of the divide." (Cran- 
brook, in Kinnear, 1934, p. 348.) 

Ratufa bicolor hainana (J. A. Allen) 

Ratufa gigantea hainana J. A. Allen, 1906, Bull, Amer. Mus., 22, p. 472. 
Ratufa gigantea stigmosa Thomas, 1923, Jour. Bombay Nat. Hist. Soc, 29, 
p. 86. 

Types. — Ratufa g. hainana, AMNH No. 26638, adult male from 
Cheteriang, Hainan, collected in 1903 or 1904 by agents of Alan 
Owstan; stigmosa, BM No., adult female from 730 me- 
ters on Doi Sritepe, Chiengmai, Siam, collected April 10, 1898, by 
T. H. Lyle. 

Material examined, from Hainan. — Mt. Wuchi (AMNH), four, 
(BM), two; Nam Fong (AMNH), one; Mts. S. W. of Kachek 
(USNM), one. 

Material examined, from Chinese mainland. — Wa-tien, Yunnan 
(AMNH), one; Wan Tien (MCZ), one; Shui-kow-kwan, Lungchow, 
Kwangsi (CNHM), one; Nam-ting River at Burma border, 1700 feet 
(AMNH), one. 

Material examined from Indochina. — Bolovens Plateau, Laos 
(AMNH), two, (BM), one; Chapa, Tonkin (MCZ), two, (BM), 
two; 150 mi. west of Xieng Khouang, Laos (MCZ), one; Mt. Fansi- 
pan. Tonkin (MCZ), one; between Harsa and Lao Fou Thai, 3000 
feet, Laos (CNHM), one; Muong Yo, 2300 feet, Laos (CNHM), 
two; Ban Phone (CNHM), two; Thateng (CNHM), two; Banteai 
(CNHM), one; Hoi Xuan, Annam (CNHM), one; Phong Saly, 4400 
feet, Laos (CNHM), one; Tam Dao, 3000 feet. Tonkin (BM), five; 
Nape [Na Pe], 2500 feet, Laos (BM), three; Phu-Qui, 100 feet, An- 
nam (BM), one, (MNHN), three; "Nghia-hung," Phu-Qui, 100 feet 
(BM), two; Xien Tuang-Koo [Xieng Khouang], Laos (BM), one; 
"Ci Norn," 1200 meters (MNHN), one. 


Material examined, from Thailand. — Doi Sutep, Chiengmai 
(AMNH), one; Doi Pui, Chiengmai (AMNH), one; Doi Hua Mot 
(USNM), two; Doi Nangka (USNM), two; Doi Nangkeo (MCZ), 
two; Mt. Angka (MCZ), three; Ching Dao (MCZ), one. 

Material examined, from Burma. — Maymyo (AMNH), one; Gok- 
teik, 2133 feet, Northern Shan States (BM), one; Katha (BM), one; 
Madaya Forest, 55 miles north of Mandalay, 2000 feet (BM), two. 

Original description. — "Whole upper parts, outsides of limbs, and 
the tail uniform intense black; ventral surface and inside of limbs 
rusty yellow, the basal half of the pelage over the chest and belly 
brownish black, showing more or less at the surface over the central 
part of the abdominal area; a broad black cheek stripe, and two small 
spots of black on chin. Ears tufted . . ." J. A. Allen gave this de- 
scription from only the one type specimen, but it is quite good for 
the additional material subsequently available from Hainan reported 
here. From our inspection of the above material, however, his sug- 
gestion that the nasal bones of hainana are longer than those of 
gigantea is not sustained. For further discussion of the characters 
of hainana, see the account of gigantea. 

Habits. — A collector and observer of this squirrel reports: "Plenti- 
ful in all big forests especially round Gokteik, quite identical in habits 
with [Indian] giant squirrels." (G. C. Shortridge in K. V. Ryley, 
1914, p. 721.) 

Genus FUNAMBULUS Lesson 

Funambulus Lesson, 1835, Illustr. de Zoologie, pi. 43, and 2 pp. of text. 
Palmista Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 279. 
Tamiodes Pocock, 1923, Proc. Zool. Soc. London, 1923, p. 215. 

Type species.— Funambulus, Sciuru^ indicus Lesson {=S. palma- 
rum Linnaeus) ; Palmista, fixed as F. palmarum Linnaeus by Thomas, 
1897, Proc. Zool, Soc. London, 1897, p. 933; Tamiodes, Sciurus tris- 
triatu^ Waterhouse. 

Definition. — The genus Funambulus comprises the species pen- 
nanti, palmarum, tristriatus, layardi, and sublineatus, all of which are 
small to medium-sized tree squirrels and endemic to the Indian Sub- 
region. (See their distributions in Figures 7-11.) 

Diagnosis. — (1) The baculum of Funambulus is a single unit, lack- 
ing the separate bony blade characteristic of some squirrels (Pocock, 
1923). (2) There is (as shown in the lateral views in Figures 5 and 6) 
no dorso-anterior process of the premaxillary bone rising to abut 
upon the anterolateral angle of the nasal bone (Moore, 1959, p. 170). 
(3) There are (as shown in the ventral views in Figures 5 and 6) one 
or two transverse bony septa crossing the auditory bulla (Moore, 
1959, p. 170). (4) There are in parous females but two pairs of func- 
tional mammae (Moore, 1961a, p. 8). (5) The dorsal pelage is longi- 
tudinally striped with 3 or 5 light stripes contrasted with brown or 
black pelage between stripes. (6) The coronoid process of the man- 
dible is low and only incipiently falcate. (Compare Figures 5 and 6 
with Figures 1, 12, 17, 22, 23, 30, and 32.) 

The above six characters distinguish Funambulus from other 
Sciurinae of the Indian and Indochinese subregions as follows: from 
Ratufa by 2, 3, 4, 5, and 6; Callosciurv^ by 1, 2, 5, and 6, Tamiops 
by 1, 2, 4, 5, and 6, Dremomys by 1, 2, 4, 5, and 6; Menetes by 1, 2, 4, 
and 5; and Sciurotamias by 2, 3, 4, 5, and 6. 

Systematic history. — For many years the genera Dremomys, La- 
riscus and Menetes were confused with Funambulus. A number of 
forms belonging in those genera were described as forms of Funam- 


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Fig. 5. Skull and left mandible of the Indian striped squirrel, genus Funam- 
bulus, species tristriatus, AMNH No. 54652, X 1. Note that there is no dorso-ante- 
rior process of the premaxillary bone rising to abut evenly with the antero-lateral 
angle of the nasal bone as is present in all other genera of Oriental squirrels. 

hulus. The relationship was clarified when Thomas (1908, 1914, 
1916a) separated Lariscus and "Zetis" (=Menetes) from Funambulus 
of the Indian peninsula and Ceylon. This distinction was further em- 
phasized by the marked differences in the bacula discovered by Po- 
cock (1923), whose consequent placement of these genera in separate 
subfamilies was recognized by Simpson (1945) as of tribal difference. 
Prasad (1951, 1954, 1957) discovered that the male reproductive tract 
of Funambulus is of a non-penile-duct type, extremely different from 
the type considered characteristic for Sciuridae, and (1957, p. 21) 
proposed that this genus be separately raised to subfamily rank. 
Moore (1959, p. 170) found a skull character which links Funambulus 
with African genera Funisciurus, Paraxerus, and Myosciurus, the 
male genital tracts of which are not known, and left these four genera 
all in the tribe Funambulini Simpson. 

Early systematic treatments of the genus Funambulus are few: 
Blyth (1849, 1852) dealt with Ceylonese races. Wroughton (1905) 
discussed the identity of F. palmarum at length and distinguished 
it from the north Indian F. pennanti. Thomas and Wroughton 
(1915b) published a key to the Funambulus of Ceylon. Wrough- 
ton (1916d) worked out the races of the species palmarum and tris- 
triatus of peninsular India. Pocock (1923) showed the characters 
of the glans penis and baculum in F. palmarum and F. tristriatus 
and solely on the differences in these separated the species tristriatus 
at the generic level as Tamiodes. Phillips (1928) discussed the vari- 


ous Funambulus from Ceylon. W. C. 0. Hill (1936) examined the 
bacula of all Ceylonese Funambulus, and was puzzled to find that 
the bacula of all the Ceylonese forms thought to belong to species 
Funambulus palmarum actually display the characters which Pocock 
(1923) ascribed to Tamiodes tristriatus and differ, just as tristriatus 
was supposed to differ, from Pocock's (1923) characterization of 
F. palmarum. Prasad (1957, p. 7) supports Hill's findings with an 
excellent figure showing the baculum of Ceylonese F. palmarum 
kelaarti to be like that of tristriatus, also. Furthermore, Prasad 
(1954, fig. 2) had already shown that the baculum of F. palmarum 
palmartim of India was also quite a good deal more like Pocock's 
figure of tristriatus than like Pocock's palmarum. It remains only, 
then, to note that Prasad's (1957, fig. 2) illustration of the baculum 
of Funambulus pennanti of northern India is like Pocock's (1923, 
fig. 20) of F. palmarum. Pocock's specimen alleged to be palmarum 
was therefore evidently pennanti. Actually, as Hill (1936) and Pra- 
sad (1954, 1957) have now shown, the bacula of Funambulus palma- 
rum indicate close relationship to F. tristriatus, and it is F. pennanti 
which is very different. This solution to the Tamiodes puzzle is 
almost equally well established by the close correspondence of the 
characters described by Pocock (1923, p. 215) for the glans penes of 
his supposed Funambulus palmarum and Tamiodes tristriatus respec- 
tively to the illustrations of F. pennanti (Prasad, 1957, fig. 1) and 
F. palmarum (Prasad, 1954, fig. 2). Furthermore, since no one has 
yet pubUshed a description or figure confirming the male genital 
characters of tristriatus, it remains possible that Pocock's (1923) 
tristriatus was in fact a specimen of palmarum and that male genital 
characters for tristriatus remain unknown.^ 

Pocock's mistake with pennanti must be a fairly easy one to make, 
for Didier (1953, p. 69, fig. 3) has evidently made it again, illustrat- 
ing in detail three views of the baculum of "Funambulus palmarum 
L. (Hindoustan) ." The figures quite evidently represent the baculum 
which Prasad (1957) has shown to be that of pennanti. 

Zahn's (1942) revision of this genus is distinguished particularly 
by his inclusion in it of the striped squirrels of the subgenus Tamiops. 
Ellerman and Morrison-Scott (1951) did not follow this superficial 
arrangement, and Moore (1959, 1961a) has produced new evidence 

1 Recognition now of the nature of the error involved in Pocock's concept of 
tristriatus as a distinct genus justifies earlier abandonment of this concept by vari- 
ous authors because of Hill's (1936) evidence of its incongruity. The difference 
recognized by Pocock (1923) still exists, differentiating pennanti (not tristriatus) 
from the other four species, and will be honored here as of subgeneric significance. 


that supports Pocock's (1923) in classifying Funambulus with cer- 
tain African squirrels in the tribe Funambulini but Tamiops with 
Oriental genera in the tribe Callosciurini. Further evidence is also 
offered in the present paper in the discussion under Tamiops. 

Parameters of the populations of subspecies of the genus Funam- 
hulus as represented by museum material would be an important 
contribution to knowledge, although it is too intensive for so exten- 
sive a revision as the present one. Some indication of the variation 
in size within the genus, however, may be seen in Table 3. The 
sample representing species palmarum in Table 3 may give a fair 
indication of where the mean for any of these measurements in that 
species might fall. 

Distribution. — This primarily tropical genus inhabits India and 
Ceylon. Three of its five species are represented by seven forms on 
Ceylon. Nine other races representing all five species, appear to 
occur on the mainland only below the Tropic of Cancer, leaving five 
races (representing only two species) which occur north of it. 

Intrageneric relationships. — The constancy of the distinctive 
striped pelage pattern causes the species of Funambulus to seem 
extremely close in relationship to one another. The most distinct 
of the five species is pennanti, for not only does it possess five white 
stripes instead of the three common to the other four species, but its 
rod-like baculum lacks any suggestion of the bifurcated tip now re- 
ported to occur in the other four species, and the shape of its glans 
penis apparently also differs from that of the other four (Hill, 1936, 
figs. 1-5; Prasad, 1954, fig. 2; Prasad, 1957, fig. 1). 

One might anticipate finding an interesting amount of difference 
between the two species of jungle squirrels, sublineatus and layardi, 
which occur in heavy jungle in southern India and Ceylon. Their 
glans penes and bacula as illustrated by Hill (1936, figs. 4 and 5) 
appear to distinguish the two from each other as much as from pal- 
marum. From a very meager amount of material, one skull of F. s. 
sublineatus, UMMZ No. 81078, three of F. s. obscurus, MCZ No. 
27550, NMC No. 60-x-E, NMC No. 60-x-C, and two of Funambulus 
I. layardi, NMC No. 59-H, and NMC No. 59-N, and a series of three 
photographic views each of the types of kathleenae (=S. s. obscurus), 
sublineatus, delesserti (= sublineatus) , and F. layardi signatus, before 
the senior author, it appears that the interorbital breadth just be- 
hind the supra-orbital notches (to avoid a broken place), is 0.81, 0.84, 
and 0.85 of the greatest length of frontals in the three layardi and is 
0.60, 0.63, and 0.68 in three of the sublineatus. If this difference is 



constant, it is a large one, and suggests an important difference in 
ecological niche. The broad frontals with more lateral orientation 
of the eyes generally characterizes squirrels whose habits take them 
high in the trees. Narrower frontals and more upward orientation 
of the eyes characterize the species which are adjusted to living close 
to, or on, the ground. Since one would expect some niche difference 
between two closely related squirrels of the same genus which are 
said to inhabit the same jungles and which differ in size, it is espe- 
cially interesting to find that Phillips (1935, pp. 238 and 241) reports 
layardi to inhabit the tops of the tallest trees in the tallest and heavi- 
est forests, but sublineatus to inhabit the "undergrowth in the jun- 
gle." See details quoted in species account below of sublineatus. 

Fig. 6. Skull and left mandible of the Indian striped squirrel genus Funam- 
bulus, species sublineatus, UMMZ No. 81078, X 1. Note the character that distin- 
guishes Funambulus from all other Oriental squirrels, absence of any dorso-anterior 
process of the premaxillary bone rising to abut evenly with the antero-lateral angle 
of the nasal bone. Note also the characters that have led to sublineatus being con- 
sidered an incipient pygmy squirrel (Moore, 1959, pp. 186-190). 

Charles McCann (letter of June 21, 1961) comments on apparent 
ecological segregation of species on the mainland nearby: "... [near] 
the Palni Mountains pennanti is found on the plains, palmarum on 
the hills up to about 4,500 feet, and above occurs the little F. subline- 
atus feeding chiefly on Rubus species in the undergrowth, its normal 
habitat. The green viper, Trimersurus macrolepis, appears to be its 
natural enemy up to 7000 feet." (This locality, latitude 10° 20' N., 
is about 5° farther south than the southernmost locality from which 
we have examined pennanti material.) 

A further aspect of the same species character which distinguishes 
the above-mentioned available material of sublineatus from layardi 
(and also from palmarum, pennanti, and tristriatu^) is obsolescence 
in sublineatus of the supra-orbital notch, which otherwise seems to 
be well developed throughout the genus. 


The material on Funambulus in American museums and hence 
available to Moore in the present study, has been extremely small 
for the assessment of the validity of subspecies and to a much lesser 
extent than for other genera has it been possible to push the study 
beyond Tate's review concept to a revision. 

Moore (1960) has pointed out that the squirrels confronting 
Funambulus pennanti across the Garo-Rajmahal Gap are tribally 
different, and that much zoogeographic importance may attach 
to detailed further field study of the distribution of F. pennanti 
and Callosciurus pygerythrus in this region, and where the two also 
appear almost to meet along the foothills of the Himalayas. See 
also a note in the account of C. pygerythrus blythi in the present 
paper. The importance and need of more data on the breeding of 
F. pennanti has also been shown (Moore, 1961, pp. 21, 27, 29). 


1. Dorsal pelage marked by five longitudinal light stripes, and baculum a simple, 

slightly curved rod pennanti 

Dorsal pelage marked by three longitudinal light stripes, and baculum bifur- 
cated at distal end 2 

2. Fur long, tending to obscure the pelage stripes, supraorbital notches absent or 

obsolescent, bifurcations of baculum ending in large knobs sublineatus 

Fur short, stripes quite distinct, supraorbital notches usually well developed; 
bifurcations of baculum without large terminal knobs 3 

3. Midstripe, at least, rather brightly colored (orange-yellow, etc.), ventral pelage 

richly colored (russet, chestnut, yellow-orange, etc.), bifurcation of baculum 

obsolescent (or only incipient?) layardi 

Midstripe and ventral pelage only pale buffy, bifurcation of baculum quite 
distinct 4 

4. Occipitonasal length usually exceeding 40 mm.; length of palate (front of in- 

cisors to back of palate) usually exceeding half of the occipitonasal length. 


Occipitonasal length usually less than 40 mm.; length of palate usually less 

than half of occipotonasal length palmarum 

PRASADSCIURUS new subgenus 

Definition. — This subgenus includes only the species Funambulus 
pennanti Wroughton which is endemic to the northern part of the 
Indian Subregion. See its range as plotted from material we exam- 
ined in Figure 7. 

Diagnosis. — The baculum is an attenuate rod without apical bi- 
furcation. The glans penis is extended apically in a slender point. 
There are five longitudinal light stripes in the pelage of the dorsum 
and sides. 


Funambulus pennanti Wroughton is the type species. The sub- 
generic name recognizes the work of the Indian scientist, Dr. M. R. N. 
Prasad, whose description of the genital characters of pennanti pro- 
vides the critical clue to clarification of the Tamiodes puzzle. 

Funambulus pennanti Wroughton 

Definition. — This species includes all of the five-striped forms of 
Funambulus and is endemic to the northern part of the Indian Sub- 
region, most of its range lying north of the Tropic of Cancer. See 
Figure 7. 

Diagnosis. — Same as for subgenus Prasadscuirus. The following 
characters seem also to have diagnostic value in the material at the 
American Museum of Natural History. The longer pair of lateral 
stripes extends posteriorly over the rump to the base of the tail, and 
clearly forward upon the shoulders and nape to reach the ear. The 
middle stripe extends posteriorly onto the tail. The head of pen- 
nanti bears two nearly parallel stripes, a faint light stripe from ear to 
eye, and a stronger, almost white one from beneath the base of the 
ear forward below the eye, and sometimes quite clearly halfway out 
to the tip of the snout from the eye. This species possesses no bright, 
warm color about the anus or extending out the ventral side of the 

Relationships to other species. — In describing this species, Wrough- 
ton (1905) remarked, "It would almost seem that palmarum is a 
South and pennanti a North Indian form . . . they occur together on 
the West Coast. . . ." Our investigation certainly supports this early 
concept of distribution of the two species. See Figures 7 and 8. 
Only F. palmarum rohertsoni and F. p. bengalensis occur within the 
range of F. pennanti, and except for a southward invasion just east 
of the Western Ghats to Dharwar (latitude 15°27'N.), the spe- 
cies pennanti seems to range only north of the twentieth parallel of 

A collector (C. A. Crump in Wroughton, 1915c, p. 108) reports 
observations on this species: "Common at Chainpur and well dis- 
tributed throughout Hazaribagh where, however, it overlaps with F. 
palmarum, the latter I think predominating. At several places I shot 
both these squirrels on the same ground but did not find them to- 
gether in any one tree." Separately one of us has treated the known 
interrelationships of these two species in more detail (Moore, 1960, 
p. 6) and speculated on their origin as species (pp. 9-12). 



Mr. Charles McCann of the Dominion Museum, Wellington, 
N. Z., contributes (letter, June, 1961) the following comments on 
pennanti from his years in India with the Bombay Natural History 
Society : 

"Habitat. — An inhabitant of the open plains and scrub country. 
It is more commensurate with man and is frequently associated with 
villages, whether on the Deccan Plain or the Sind Desert. Does not 
occur in jungle country except, perhaps, on the outskirts. 

"Coat. — Harsher, smoother, and lying more flat than in palmarum. 

B • 



y 'N. 


• • 




1 ' I I I 



4- +I6°N 

Fig. 7. Geographical distribution of the five-striped Indian squirrel, Funam- 
bulus pennanti, as revealed by specimens examined, except that one record of 
argentescens is about 275 miles off the map at Mand, Baluchistan. Subspecies: 
A, pennanti; B, argentescens; C, lutescens. 


"Voice. — Whistle-like, shriller, sharper and more prolonged than 
that of palmarum. 

"Nest.— Builds nest under tiles and eaves of buildings and in 
hollow trunks of trees and among palm leaf sheaths. In the desert, 
owing to the absence of suitable nesting sites, it builds like palmarum, 
but the nest is globular and more disorderly. It will sometimes com- 
mandeer the nest of Uroloncha. Coconut fibre is in great demand 
when available — hence, its nest-building is destructive to mattresses 
placed out for airing. 

"Habit. — More terrestrial than palmarum. Perhaps more om- 
nivorous than palmarum." 

Funambulus pennanti pennanti Wroughton 

Funambulus pennanti Wroughton, 1905, Jour. Bombay Nat. Hist. Soc, 16, 
p. 411. 

Type.— BM No., adult female from Mandvi Taluka, 
Bundha, Surat District, India, collected February 27, 1898, by 
R. C. Wroughton. 

Material examined, from Bombay. — Bombay (BM), three; Shen- 
durni, 900 feet (BM), one; Poona, 1900 feet, Deccan (BM), five; 
Karad, Satara District (BM), one; Nasith [Nasik] (BM), one; 
"Mehda," Satara District (BM), one; "Satapur " 20 feet, Dhranga- 
dhra ^BM), four; Dharwar, South Mahratta (BM), one; Bodbad 
[Bodvad], 1000 feet (BM), one; Parola, 880 feet (BM) two. 

Material examined, from Berar. — Akola (UMMZ), one. 

Material examined, from Central Provinces. — Balaghat District 
(CNHM), three; "PhichpalH," 1300 feet, Chanda (BM), two; 
"Thain," 1400 feet, Hoshangahad (BM), two; "Ganoor," 1000 feet, 
Nimar (BM), one; "Sakot," 1200 feet, Hoshangahad (BM), one; 
"Hewra," 1000 feet, Nimar (BM), two; Sohajpur, 1000 feet, Hoshan- 
gahad (BM), one; Chanda, 800 feet (BM), two; "Rithi, C. P." 
(AMNH), one; Khandwa (AMNH), two; Raipur (AMNH), five; 
Asirgarh, 1500 feet, Nimar (BM), one. 

Material examined, from Gujerat. — Danta, 1000 feet (BM), one. 

Material examined, from Kathiawar. — Junagadh [Junagarh], 480 
feet (BM), two; Rajkot, 100 feet (BM), two; "Sadla," 10 feet, Bajana 
State (BM), two; Sehore [Sihor], 1600 feet, Bhaonagar Province 
(BM), four; Talala [Talaja], 200 feet, Junagarh State (BM), two; 
Van Kaneer [Vankaner], 500 feet (BM), three. 


Material examined, from Sind, West Pakistan. — "Pano Agil," 
Upper Sind (BM), five; Shikarpore [Shikarpur] (BM), one; Sehwan 
(BM), one. 

Material examined, from Rajputana. — Mt. Abu, 4300 feet (BM), 
one; Sambhar (BM), one. 

Material examined, from Punjab. — "Azadpur," 550 feet, Delhi 
(BM), one; Lahore (UMMZ), one; Sirsa, Hissar (UMMZ), five; 
"Bhadwar," Kangra (UMMZ), three; "Khajjean" Kangra (UMMZ), 

Material examined, from United Provinces. — Pihbhit, 800 feet, 
Rohilkand (BM), two; "Dela," 1500 feet, Ramnagar, Kumaon (BM), 
one; "Dachawri," 2500 feet, Kumaon (BM), one; "Jerna," 1500 feet, 
Ramnagar (BM), one; Ramnagar, 1100 feet, Kumaon (BM), seven; 
SiwaUk Hills (BM), one; Etawah (UMMZ), two. 

Material examined, from GwaHor. — Bhind (BM), three; Corepura 
[Chorepura], 1100 feet (BM), two; "Ghatigoan," 900 feet (BM), 
four; Morar (BM), one. 

Material examined, from Nepal. — Tribeni [Ghat], Terai (BM), 
four; Hetora [Hataura] (AMNH), one; "Pili," Sipora or Sipna [Sipra] 
Valley (BM), one; "Bandasa," West Terai (BM), two. 

Material examined, from Bihar. — "Jagodih," 600 feet, Hadari- 
dagh [Hazaribagh] (BM), two; Daltonganj, 600 feet (BM), one. 

Material examined, from Bengal, East Pakistan. — Chandpara 
(AMNH), three; Salbani, 200 feet, Midnapore (BM), two; Sevoke 
(CNHM), one; Kharagpur (UMMZ), one. 

Material examined, from Cooch Behar. — Haldibari (MCZ), one. 

Original description. — ". . . body colouring is very much as in 
palmarum, comorinus, but along the outside edge of the 'saddle mark' 
on each side there is a supplementary pale stripe . . . bounded on the 
outer side by the general body colour . . . where it is commencing to 
pale down to the meeting line with the belly colour. . . . No band 
of short, rufous hairs along the midrib under the tail as there is in 
palmarum. ..." 

Funambulus pennanti lutescens (Wroughton) 

Funambulus lutescens Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, 
p. 429. 

Type.— BM No., adult female from Deesa, 450 feet, 
Palanpur, Gugerat, India, collected April 29, 1913, by C. A. Crump. 


Original description. — ". . . rather smaller than true pennanti, 
about the size of argentescens, but much paler than either of these 
two forms. . . . General colour above pale 'cream buff' on the flanks, 
the saddle near 'mars brown' . . . the five longitudinal stripes white, 
very slightly tinged with buff, the lateral pair scarcely distinguish- 
able. . . . Face drab. Hands and feet buffy white. . . . Below white, 
the hairs white to their bases. Tail also pale buffy white, each hair 
with two black rings. . . . 

"This form ranges south as far as the northern part of Kathiawar, 
but thereafter the specimens grow darker . . . and show a passage to 
true pennanti." 

In lutescens the lateral stripe runs undiminished from the ear to 
the hind limb; its width 6 mm. 

Funambulus pennanti argentescens Wroughton 

Funambulus pennanti argentescens Wroughton, 1905, Jour. Bombay Nat. Hist. 
Soc, 16, p. 412. 

Type.— BM No., adult male from Rawalpindi, 1670 feet, 
Punjab, Pakistan, collected December 10, 1900, by Birrell. 

Material examined.- — Mand, Baluchistan (BM), one; Gajar, 3200 
feet, Mashkai, Kalat (BM), two; Bohara [Bahara], Sind (BM), one; 
Garo [Gharo], Sind (BM), one; Jacobabad, Sind (BM), two; Kash- 
mor, North Sind (BM), two; Mirpur Sakro, Sind (BM), three; Jhim- 
pir Lake, Sind (UMMZ), two; "Khinjar Lake," Sind (UMMZ), 
three; Karachi, Sind (AMNH), one; Kohat, 1700 feet. North West 
Frontier Province (BM), three; Peshawar, 1700 feet (BM), two; 
Amballa [Ambala], Punjab (MCZ), six, (BM), one; "Ara," 2100 feet. 
Salt Range (BM), two; "Choan," 2400 feet. Salt Range (BM), one; 
Kallak Kahar [Kallar Kahar], 2113 feet. Salt Range (BM), two; 
Multan, 600 feet (BM), four; Rawal Pindi [Rawalpindi], 1671 feet 
(BM), 10; "Kotla," Kangra District (AMNH), six; Sodhi, 2000 
feet. Salt Range (BM), one; Gholam (BM), one; Turbat, 600 feet, 
Kech (BM), three; Malasa, United Provinces (AMNH), one; "Lar- 
kanat," Naundero [Nandero] (BM), two; Kooloo Valley [Kulu Val- 
ley] (MCZ), two; Chak-Lala [Chakwal] 1100 feet, Rawalpindi, N. W. 
Punjab (CNHM), two. 

Original description. — ". . . Pattern . . . identical . . . with . . . 
pennanti . . . much paler, however, and almost all rufous tint has dis- 
appeared, . . . body colour is a pale French Gray . . . stripes and belly 
bright white. . . ." 


The pelage is very short. The three white dorsal stripes are each 
5 mm. wide. Dorsal pelage is dark reddish brown, grizzled, becom- 
ing pale gray on the shoulders, neck, and head. The limbs are pale 
gray. Ventral pelage is white the full length of the hairs. 

This subspecies is distinguished from F. pennanti pennanti by 
whiter tail, paler feet (Pale Olive Buff), and paler body color. That 
the races of F. pennanti can apparently be distinguished by the num- 
ber of black bands on the tail hairs was noted too late for testing on 
all the material examined, but we find that it separates the above 
five lots of p. pennanti from the two of p. argentescens of the AMNH 
material; two bands in p. pennanti, one in p. argentescens. 

Funambulus palmarum (Linnaeus) 

Definition. — This species includes those three-striped squirrels of 
the Indian Subregion, distributions of which are shown by collecting 
localities for the specimens we have examined, in Figure 8. (See also 
the synonyms listed in the subspecies accounts.) 

Diagnosis. — (1) The baculum is distally bifurcated or makes a 
right angle turn near the distal end, but the apex or apices are not 
knobbed. (2) The dorsal pelage is marked by three very distinct 
longitudinal hght stripes. (3) The occipitonasal length is usually 
less than 40 mm. (4) Measurement from front of incisor to back of 
palate is usually less than half the occipitonasal length. (5) The pel- 
age is relatively short and harsh. (6) The midstripe and ventral 
pelage are no more richly colored than pale buffy. 

The above characters distinguish palmarum from the other con- 
generic species as follows: pennanti by 1 and 2; tristriatus by 3 and 4; 
layardi by 6; and sublineatus by 1, 2, and 5. 

Relationship to other species. — Mr. Charles McCann of the Do- 
mion Museum, Wellington, N. Z., contributes the following remarks 
(letter of June, 1961) on F. palmarum from his years with the Bom- 
bay Natural History Society in India: 

"Habitat. — More a hill forest species than is pennanti, inhabiting 
deciduous rain forest. Ascends to about 4,000-4,500 ft. In such 
localities it will approach human environments, taking the place of 
pennanti. On the Panchgani-Mahableshwar plateau, 4100 ft., (out- 
skirts of the Western Ghats), palmarum occurs on the hills, but is 
replaced by pennanti on the Deccan Plain (= 1800 ft.). F. palmarum 
occurs almost throughout the Western Ghats in deciduous rain forest 
but does not enter the wet evergreen forest. 


• D 






• • C 






20°N + 





y"^ +18° N 

/ 86° 



, • S • ,•■■■" 

+ ^ 

1 • 

\ +/4°N 
\ 82» 

\ • • 


I2°N + 

\ • • / 

/ + 


BS^ 82° 

8°N+ ^^ \ 
76° 78° \ 

^ ^ \CEYLON 



MILES 400 
1 < 1 


Fig. 8. Species range of the Indian palm squirrel, Funambulus palmarum, as 
shown by specimens examined. Subspecies: A, palmarum; B, bellaricus; C, robert- 
soni; D, bengalensis; E, comorinus; F, favonicus; G, kelaarti; H, brodiei, and 
I, olympitis. The combined ranges of species palmarum and pennanti approximate 
the extent of the Indian Subregion. 



"Coat. — Soft and somewhat silky and in life more erect than that 
of pennanti. 

"Voice. — More bird -like and slightly deeper than that of pennanti. 

"Nest. — Builds nest like a passerine bird in branches, somewhat 

"Habit. — More arboreal but does come down to the ground. Nec- 
tar is one of its foods. Bark searching is a frequent habit, possibly 
for insects. Fruit appears to be its main diet." 

Funambulus palmarum palmarum (Linnaeus) 

Sciurus palmarum Linnaeus, 1766, Systema Naturae, 12th ed., 1, p. 86. 
Sciurtis penicillatus Leach, 1814, Zool. Misc., 1, p. 6, pi. 1. 
Sciurus indicus Lesson, 1835, Illustr. de Zoologie, no. 15, pi. 43, 2 pp. text. 
Funambulus gossei Wroughton and Davidson, 1919, Jour. Bombay Nat. Hist. 
Soc, 26, p. 730. 

Types. — Sciurus palmarum, apparently lost, not found at Upsala, 
locality restricted to vicinity of Madras, India (Wroughton, 1905, 
p. 410); penicillatus, not found; indicus, not found; gossei (BM) No., an adult male from Kotagiri, 4100-4500 feet, Nilgiri Hills, 
taken June 20, 1918, by PhiHp Gosse. 

Material examined, from Madras Province. — "Adyar" [in city of 
Madras] (BM), two; "Kilcauk" [Kilpauk in city of Madras] (BM), 
two; "Museum Grounds" [city of] Madras (BM), two; "Panampet" 
(BM), one; "Tadiarpet" (BM), one; Trichinopoli, 400 feet (BM), 
two; Chittoor (AMNH), one; Salem District (AMNH), two. 

Material examined, from other provinces. — Coimbatore (BM), 
one, (CNHM), one; Polkonda Hills, 1000 feet, S. Cuddapah (BM), 
two; Shevaroy Hills, 4500 feet, E. Ghats (BM), one; "Sweat Dist." 
(BM), one; "Machen," 4000 feet (BM), one; Kanara at S. Mahratta 
border, 2000 feet (BM), one; Seringapatam, 2338 feet, S. Mysore 
(BM), three; Kolar, 2786-4026 feet, E. Mysore (BM), two; Kurum- 
dapatti, Salem Distr. (BM), three; Kalhatti [Kalpatta], Nilgiri Hills 
(AMNH), one; "Kellengode" (AMNH), one; "Honnametti Estate," 
Biligirirangan Hills, Mysore (AMNH), one; Anaikatti, Nilgiri Hills 
(AMNH), one; "Gantha," 1300 feet, Coimbatore, southern India 
(BM), one; "Rookery Kil," 4500 feet, Kotagiri, Nilgiris (BM), three. 

Although gossei was originally described as belonging to the "tri- 
striatus group," Ellerman (1940, p. 379) held it a separate species 
by itself, and Zahn (1942, p. 58) placed it in the species palmarum. 


Comparison of the dimensions of the type of gossei in our Table 3 
with those of species palmarum and species tristriatus supports Zahn's 
allocation. Ellerman and Morrison-Scott (1951, p. 494) synonymize 
gossei under F. p. palmarum, and this seems to us the most reason- 
able treatment. 

The color of F. p. palmarum, as evidenced by the above series of 
three from the Salem District and Chittoor near the type locality: 
A middorsal area Mummy Brown to Prout's Brown pales anteriorly 
behind the shoulders and posteriorly on the rump. Through this 
run three longitudinal lines of Light Buff to Light Ochraceous Buff 
about 3 to 4 mm. wide. The middle one of these extends from the 
nape (into which it may extend faintly) to the base of the tail. The 
lateral ones separated by 11 to 12 mm. from the middle stripe, extend 
from back of the shoulders onto the rump, failing to match the length 
of the middle stripe by about 10 mm. anteriorly and perhaps 3 to 
5 mm. posteriorly. Five to 7 mm. below the lateral lines, the 
Mummy Brown dorsal area changes abruptly to the agouti sides, 
which are Smoke Gray to Apricot Buff. The color of the sides ex- 
tends onto the shoulders, nape, cheeks, ears, and rostrum, and onto 
the flanks and all limbs and feet. The crown color is intermediate 
between that of the sides and back. There is an almost white area 
about 2 by 10 mm. just medial to each ear, oriented anteroposteri- 
orly, and fading into the body color posteriorly. There is a light- 
colored "eye-ring" 1 mm. wide above and below the eye, but not 
enclosing it at either end. The tail is somewhat less than half the 
animal's total length, about 20 mm. wide. Its dorsal and lateral hairs 
are whitish at the base, middle, and tip, with two black bands be- 
tween and these give the tail as a whole an appearance of jumbled 
annulation (with about 15 black rings in one instance). On the hairs 
of the lower surface of the tail the basal and middle portions of each 
hair are Cinnamon-Rufous proximally on the tail grading out to 
Apricot Buff at its end. This Cinnamon-Rufous color extends onto 
the ventral pelage of the body in a small area about the anus. The 
ventral pelage from this to the chin and on the under sides of all the 
legs is white with faint washings of Maize Yellow. 

Habits. — G. C. Shortridge remarks of Funambulus palmarum col- 
lected in the vicinity of Dharwar (in Wroughton, 1912, p. 1186), 
"They may often be found in prickly pear thickets when their mouths 
are usually stained crimson with the juice of the fruit. Both this 
species and tristriatus feed also on the berries of the Lantana." 


Funambulus palmarum comorinus Wroughton 

Funambulus palmarum comorinus Wroughton, 1905, Jour. Bombay Nat. Hist. 
Sec, 16, p. 411. 

Type.— BM No., adult female from Katyani, Trevan- 
drum, Travancore, India, collected January 23, 1895, by H. Ferguson. 

Material examined. — Nagercoil (BM); one, "Benhope," 3000- 
4000 feet, Nilgiris (BM), three; Kotagiri, Nilgiris (BM), two; Tre- 
vandrum, Trevancore [Travancore] (BM), 11. 

Original description. — "Differs from [palmarum] by its much so- 
berer grey colouration and larger skull measurements." 

This race, generally much darker in color than the true palma- 
rum, is evidently closely related to the dark races of Ceylon. The 
width of the median line is between 2 and 3 mm. The brown of the 
dorsal colored area between each lateral line is considerably narrower 
than in other races of palmarum. The hairs of the under parts are 
white or dirty white, with short gray bases. The anal area is brown, 
as in other Indian forms. 

Funambulus palmarum bellaricus Wroughton 

Funambulus palmarum bellaricus Wroughton, 1916, Jour. Bombay Nat. Hist. 
Soc, 24, p. 647. 

Type. — BM No., adult male from Vizayanajar, Bellary, 
India, collected July 20, 1912, by G. C. Shortridge. 

Material examined. — Vizayanajar, 1500 feet, Bellary, India ''CN- 
HM), two. 

Original description. — ". . . differs [from palmarum] in the com- 
plete absence of yellow suffusion on the forearms, shoulders, and 
thighs. General colour above rather a coarse grizzle of black and 
white, giving the effect of pale 'smoke grey.' The whole dorsal area 
forming *a saddle,' which is coloured much darker. . . . The usual 
three longitudinal dorsal stripes creamy white. The face more or 
less suffused with ochraceous, according to season. Feet and hands 
pale. . . . Tail black and white, very indistinctly barred, and below 
'orange rufous.' 

"Shortridge obtained ... 29 specimens in s. Mysore . . . consid- 
ered intermediates [between bellaricus and palmarum]." 

This subspecies is sharply separated from subspecies palmarum 
by the much greater width of the white median line, 6 mm. com- 
pared to 3. 


Funambulus palmarum robertsoni Wroughton 

Funambulus palmarum robertsoni Wroughton, 1916, Jour. Bombay Nat. Hist. 
Soc, 24, p. 647. 

Type.— BM No., adult male from Pachmarhi, Ho- 
shangabad, India, collected March 20, 1912, by C. A. Crump. 

Material examined. — Pachmarhi, 3300 feet, Hoshangbad (CNHM), 
two; Asirgarh, 1500 feet, Nimar (BM), one; Kol Kaz, 1600 feet, 
Berar (BM), one; Rarighat, 2500 feet, Hoshangabad (BM), one; 
Siwal, 1000 feet, Nimar (BM), one; Pachmarhi, 3300 feet, Ho- 
shangabad (BM), one; "Thain," 2000 feet, Hoshangabad (BM), two; 
"Nawapara," Raipur, Central Provinces (AMNH), one. 

Original description. — ". . . Sombre coloured . . . markedly smaller 
than palmarum . . . saddle brown, with a slight yellow tinge . . . get- 
ting darker at certain seasons. The dorsal stripes buffy white. . . . 
Below, in most cases, 'vinaceous cinnamon,' in some dull white. . . ." 

This subspecies differs from others in having no brown on the 
head; the gray of the sides and nape continues forward onto the 
crown and face. The width of the median line is nearly five milli- 

Funambulus palmarum bengalensis Wroughton 

Funambulus palmarum bengalensis Wroughton, 1916, Jour. Bombay Nat. Hist. 
Soc, 24, p. 648. 

Type.— BM No., adult female from Gayhundi, 1000 
feet, Hazaribagh, Bihar, India, collected May 10, 1914, by C. A. 

Material examined. — "Jagodih," 600 feet, Hazaribagh, Bihar and 
Orissa (BM), one; Muhammedganj, Behar (AMNH), one. 

Original description. — ". . . resembling robertsoni in its small size, 
but distinguishable by its much larger teeth and ... an ochraceous 
tinge on flank. Below invariably dull white." 

The width of the median line is about three millimeters. 

The pelage of the example from Muhammedganj differs from our 
description of the typical race: in having a Blackish Brown (2) dor- 
sal area; in having the stripes extend, albeit faintly, upon the shoul- 
ders to the nape; in having the mid-stripe but half the width of the 
laterals; in having the Cinnamon -Rufous of the anal area extend out 
the ventral pelage of the tail about 20 mm. before paling to Light 
Ochraceous Buff, then Light Buff; and in having a fuller tail (30 mm. 
wide with hairs to 25 mm. long). 


Habits. — A field observer collecting this form makes the follow- 
ing remarks: "Blanford states this squirrel is not found in forests, 
but in my experience though partial to the neighbourhood of culti- 
vation, it may like pennanti be found far into the forests." (C. A. 
Crump in Wroughton, 1915c, p. 108.) 

Funambulus palmarum favonicus Thomas and Wroughton 

Funambulus palmarum favonicus Thomas and Wroughton, 1915, Jour. Bom- 
bay Nat. Hist. Soc, 24, p. 39. 

Funambulus palmxirum matugamensis Lindsay, 1926, Jour. Bombay Nat. Hist. 
Soc, 31, p. 239. 

Types. — Funambulus p. favonicus, BM No., young (orig- 
inal description says "adult") female from Udugama, Southern 
Province, Ceylon, collected April 23, 1913, by E. W. Mayor; F. p. 
matugamensis, BM No., adult female from Anisigalla, 
Matugama, 30 miles southeast of Columbo, Ceylon. 

Material examined. — "Anesagalla" [Anisigalla] 100 feet, Matu- 
gama, Kalutara, Western Province (BM), 18; "St. George," 300 
feet, Matugama, Western Province (BM), one; Kottawa, Southern 
Province (BM), two; Ranna, Southern Province (BM), one; Udu- 
gama, Southern Province (BM), six (topotypes). 

Original description. — ". . . darker than true palmarum, owing to 
the greater amount of black grizzling. The saddle much darker. . . . 
The central dorsal stripe white, the lateral ones ochraceous buff. 
Below white. Tail below 'cinnamon rufous' . . . i.e. red rather than 
yellow. Feet a fine grizzle of black and buff. . . . When these [18] 
specimens were laid out in a row their warmer colouring, strong col- 
our contrasts, and grizzled feet differentiate them from the sober, 
dull-coloured palmarum palmarum." 

The top of the face and rostrum of the type is a rather bright 
reddish brown. The under parts are dull whitish with gray hair 
bases. The under side of the tail is yellowish-brown. 

Dimensions. — Phillips (1935, p. 231) provides averages of mea- 
urements in millimeters for 30 males and 30 females of favonicus. 
These are respectively: length of head and body, 144.7 and 142; 
length of tail, 124.4 and 137; length of hind foot, 34.5 and 34.5; length 
of ear, 15.2 and 16. He comments that many had the ends of their 
tails "damaged," especially the males. We take this to mean that 
the tails of many were short because the end was missing. He also 
gives maxima for these measurements. These are, for males and 
females respectively: 180 and 150, 157 and 169, 37 and 37, 17 and 17. 


Funambulus palmarum kelaarti (Blyth) 

Sciurus kelaarti Blyth, 1851, Jour. Asiatic Soc. Bengal, 20, p. 166. 

Type. — IM 9479, a parous female from Hambantotte, Ceylon, 
collected in 1845 by E. L. Layard. 

Material examined, all from Ceylon. — Aripo [Arippu], northwest 
Ceylon^ (BM), two; A'pura [Anuradhapura], North Central Prov- 
ince (BM), one; Kala-oya [a river]. North Western Province (BM), 
nine; Putlam [Puttalam], North Western Province (BM), one; Man- 
keni, Eastern Province (BM), two; Inganiyagala, 100 feet, near Kal- 
munai, Eastern Province (BM), five; Hambanpota [Hambantota], 
Southern Province (BM), 11; Wellawaya, 608 feet, Uva Province 
(BM), two; "Micouralia" (BM), one. 

Original description. — ". . . kelaarti entirely replaces all the other 
small Sciuri from Tangalle and Hambantotte, and I fancy extends 
round to Trincomali. ... It [is] . . . like . . . palmarum of India, but 
the head is much redder, the halves of the back and belly are more 
blended, and the animal is altogether smaller. . . ." 

Previously, Blyth (1849, p. 602) had written substantially the 
above in a footnote, without, however, giving the squirrel in ques- 
tion a name. 

Dimensions. — Phillips (1935, p. 229) presents averages of meas- 
urements in millimeters taken on eight males and six females of 
kelaarti. These are respectively: length of head and body, 143 and 
139.6; length of tail, 136.7 and 106; length of hind foot, 34.5 and 35.5; 
length of ear, 18 and 17.5. (Note that Phillips' average measure- 
ments of the tails of favonicus evidently included ones with part of 
their natural length missing.) He also gave maxima for these dimen- 
sions, and these are respectively: 149 and 151, 151 and 160, 37 and 
37, 20 and 19. 

Type description.- — Color characteristics of the type of kelaarti, 
examined in 1960: The pelage of the crown is Russet (XV) and ends 
abruptly along a line between the ears. The agouti pelage of the 
nape and shoulders is close to Dresden Brown, or, if one ignore the 
yellowing effect of the light bands, Saccardo's Umber (XXIV). The 
postauricular patch is Pinkish Buff. The eye ring and sides of face 
below and in front of the eye are Light Ochraceous Buff (XV) . The 
pelage color approaches Ochraceous Tawny between the eye and ear. 

' There is another Arippu in Eastern Province (latitude 08° 19' N., longitude 
81° 20' E.) which may better be the source of this material. So treated on dis- 
tribution map. 


Dark pelage on the back between the Unes is Mummy Brown at its 
greatest intensity at the middle of its length, gradually grading into 
the shoulder color anteriorly, but going through the redder Prouts 
Brown and then Mars Brown before grading into the Dresden Brown 
of the rump and thighs. The dark saddle area outside of the light 
lines borders abruptly on the sides without diminution of intensity. 
The sides are about Isabella Color (XXX). The dorsal pelage on 
the limbs is like that of the shoulders but paler. The three light lines 
on the back are Light Ochraceous Buff, the lateral ones reach five 
millimeters wide, the middle one only two. The light lines disappear 
in the shoulder and rump areas but are intense for 70 or 80 milli- 
meters. The ventral pelage is Naples Yellow (XVI) and borders on 
the color of the sides rather abruptly. The midvane of the under 
side of the tail is Tawny (XV). The tail hairs have three blackish 
bands, the proximal two each about 2 mm. long, the distal one about 
4 mm. The tail pelage is worn and a little faded but a black pencil 
is evident at the tip, and the hairs are about 30 mm. long. 

Funambulus palmarum brodiei (Blyth) 

Sciurus brodiei Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 602. 
Funambulus palmarum brodiei, Thomas and Wroughton, 1915, Jour. Bombay 
Nat. Hist. Soc, 24, p. 40. 

Type. — IM No. 9840, collected by E. L. Layard in Ceylon. 

Material examined. — Cheddikakulam [Cheddikulam], Northern 
Province, Ceylon (BM), nine, (CNHM), two. 

Original description. — "Very similar to [tristriatus] but distin- 
guished by its considerably paler colour, and especially by having 
a very long pencil tuft {S}/2 in.) at the extremity of tail, quite differ- 
ent from what is ever seen in tristriatus . . . 'confined to Palmyra-tree 
district from Puttulam to Jaffna. How much further round the coast 
I do not know." 

Type description. — The type of Funambulus brodiei has the gen- 
eral appearance of palmarum. The three dorsal stripes, however, 
pass through a dark saddle-like area on the back, about 60 mm. long 
and 40 mm. in overall width. The width of the dark pelage between 
median and lateral pale lines is 9 mm., its width beyond the lateral 
lines about 6 mm. The pale median stripe is 4 mm. wide, the lat- 
eral ones 6 mm. The dorsal color becomes russet on the head and 
rostrum. The tail is a grizzle of black and buff. The hands and feet 
are light grayish buff. The under parts to the chin, and up the sides 


of the head to the bases of the ears, also the insides of the Hmbs, are 
dull buffy yellow. The skull of the type is very badly broken and 
the nasals missing. The teeth show a moderate degree of wear, but 
the almost unworn condition of the fourth premolar and first molar 
suggests that the animal is a young adult. The maxilla spreads over 
a considerable area on the dorsal surface of the skull. 

Dimensions. — Phillips (1935, p. 227) records averages of measure- 
ments made in millimeters on but four males and three females of 
brodiei, and these are respectively: length of head and body, 150 and 
148.7; length of tail, 148 and 139; length of hind foot, 35 and 34; 
and length of ear, 17 and 14.3. The maxima that he provides with 
the above dimensions are respectively: 162 and 153, 148 and 146, 
37 and 34, 17 and 15. 

Funambulus palmarum olympius Thomas and Wroughton 

Funambulus palmarum olympius Thomas and Wroughton, 1915, Jour. Bom- 
bay Nat. Hist. Soc, 24, p. 41. 

Type.— BM No., adult female from Urugalla, 1600 feet 
central highlands, Ceylon, collected February 25, 1914, by E. W. 

Material examined. — Kandy (BM), one; "Lindoehn," 4200 feet 
(BM), one; Perodeniya [Peradeniya], 1600 feet (BM), 11; "Urugala," 
1600 feet, Central Province (BM), nine (topotypes), (CNHM), two. 

Original description. — "A dark highland form. ResembHng bro- 
diei but much darker. The saddle is commonly almost black and 
the central dorsal stripe paler than the lateral ones, often white. 
Lower side of the midrib of tail is much darker chestnut than in 
brodiei. Feet are darker even than in favonicus. Below dull white." 

This is quite the darkest of the races of palmarum; the general 
dorsal color is blackish gray. The face is brownish-gray — about like 
the sides of the body. The under parts are dull whitish with gray 
hair bases. The under side of the tail is yellow-brown. 

Dimensions. — Phillips (1935, p. 234) provides averages and max- 
ima of measurements taken in millimeters on 13 males and 11 females, 
and the averages are respectively: length of head and body, 157.5 
and 156.5; length of tail, 150.5 and 135.5; length of hind foot, 38.5 and 
37.9; length of ear, 17.5 and 17.5. The maxima are: 166 and 164, 
length of head and body, 168 and 167, length of tail, 40 and 40, 
length of hind foot, and 20 and 20 length of ear. 


Funambulus tristriatus (Waterhouse) 

Definition. — This species is constituted by those large, dark, three- 
striped forms inhabiting the evergreen rainforest of the Western 
Ghats of peninsular India as shown in our map Figure 9. 

Diagnosis. — (1) Funambulus tristriatus has three light longitudi- 
nal lines in the dorsal pelage. (2) The occipitonasal length of skull 
in adults with tooth wear exceeds 40 mm. 

The above characters distinguish species tristriatus from the con- 
generic species as follows: pennanti by 1 ; palmarum by 2; layardi by 2; 
and suhlineatus by 2. 

Systematic history. — The distinctness of Funambulus tristriatus 
from palmarum was recognized first by Waterhouse, and subse- 
quently by Wroughton (1905, p. 410), as "the forest form of palma- 
rum." Later Wroughton (1916d) distinguished the two species by 
the larger skulls of tristriatus, and in the same article described sev- 
eral new races. Later Wroughton and Davidson (1919) proposed 
two additional forms, one of which extended the range of the species 
northward along the west coast to Bombay. Pocock (1923) then (in 
error as shown above) distinguished this tristriatus group generically 
as Tamiodes. 

Relationships to other species. — Funambulus tristriatus is consid- 
erably larger than either palmarum or pennanti (see Table 3) and 
is much darker, except for the dark race of palmarum on Ceylon. 
The tail is much fuller — its width across the pelage about 25 mm. 
The longitudinal pale lines are only three. The median line is weak, 
dull yellow and 3 mm. wide; the two lateral lines are dull white, and 
about 4 mm. wide. The under parts are dull buffy white, with dis- 
tinctly gray bases to the hairs. The scrotal area and under side of 
tail (bases of tail hairs) are russet. 

This purely tropical species occurs in the forests of the mountains 
along the western coast of the Indian peninsula from about latitude 
19° N., or the city of Bombay, southward well into Travancore. 
Study to define further the distribution of subspecies tristriatus and 
wroughtoni in relation to each other and to species palmarum in 
Coorg, Mysore, Malabar, and the Nilgiris would be rewarding. 

Funambulus tristriatus tristriatus (Waterhouse) 

Sciurus tristriatus Waterhouse, 1837, Mag. Nat. Hist., (new ser.), 1, p. 499. 
Sciurus dussumieri Milne-Edwards, 1867, Rev. Mag. Zool., 19, p. 226. 
Funambulus tristriatus annandalei Robinson, 1917, Rec. Indian Mus., 13, p. 41. 

Fig. 9, Ranges of the geographic races of the Western Ghats squirrel, Funam- 
bulus tristriatus, as shown by material examined. Subspecies: A, tristriatus; 
B, wroughtoni; C, numaris. 



Types. — Sciurus tristriatus, BM No., adult male 
from "India," here restricted to the Western Ghats south of 12° N. 
latitude; Sciurus du^sumieri, MNHN No. 1838 (292), adult from 
Malabar; Funambulus t. annandalei (not seen), IM No. 8498, adult 
female, from Sasthancotta, west side of Western Ghats, Travancore, 
collected November 8, 1908, by N. Annandale. 

Material examined. — "Paumpa," North Travancore (BM), three; 
"Merchiston," Travancore (BM), two; "Paibuna," North Travan- 
core (BM), one; "Poothota," Vycoona District, North Travancore 
(BM), three; "Travancore," 4000 feet (BM), six; "Tyebautuchary," 
North Travancore (BM), two; "Bonaccord," Glen Brith Estate (BM), 
one; "India" (MCZ), one; "Kellengode," South India (AMNH), 
two; Kuttyani [Kuttyadi, Malabar], 250 feet (BM), one; "Mella- 
cotta," Wynaad (BM), one; "Madras" (BM), five; "Poumodi" [Pon- 
mudi, Travancore], 2000 feet (BM), two. 

Width of the lateral line is 4 mm., of the median line 2 mm. in 
the type specimen. The type of dussumieri has the median pale line 
45 mm. in length, and the lateral lines 70 mm. The widths are: 
median line, 2 mm.; laterals, between 3 and 4 mm. The under side 
of the tail is rather bright rufescent from the vent almost to the tip. 

Discussion. — Miss Ryley (1913, p. 437) remarked that the type 
of tristriatus "... very probably . . . came from Travancore as it 
agrees best with a small series from that district." Nevertheless, 
when Robinson (1917, p. 41) described annandalei from Travancore, 
citing this paper of Miss Ryley, he ignored the above-quoted obser- 
vation of hers and commented, "In default of authenticated skins 
from Madras I have taken modern skins from Kanara as typical of 
F. . . . tristriatus, Waterh., though it is by no means impossible that 
these will prove to represent yet another form." The Kanara series 
had in fact already been identified as belonging to a new subspecies 
of tristriatus described by Wroughton (1916, p. 646), who tersely re- 
stricted the type locality of tristriatus to Travancore. Wroughton 
and Davidson (1919, p. 728) then reported, "The British Museum 
has a series, sent by Capt. H. Ferguson, from Trevandrum [in Trav- 
ancore], which are undoubted tristriatus, with the type of which they 
agree in all essential particulars." Zahn (1942, p. 68) records the 
type locality of tristriatus as Travancore. EUerman (1940, p. 379) 
and Ellerman and Morrison-Scott (1951, p. 495) employ "Madras, 
India (by designation)." The use of "Madras" is unfortunate, for 
although the state of Madras formerly included what was still earlier 
Travancore, it does not now, and in any case Madras includes a 


much broader area than that in which anyone presumes subspecies 
tristriatus to occur. The correct political designation at the time of 
our writing is southern Kerala, but Western Ghats south of 12° north 
latitude is a better description, and in using this we are honoring and 
slightly refining Wroughton's (1916, p. 645) restriction of the type 
locality to Travancore. 

Funambulus tristriatus wroughtoni Ryley 

Funambulus wroughtoni Ryley, 1913, Jour. Bombay Nat. Hist. Soc, 22, p. 437. 

Types.— F. wroughtoni, BM No., old female from Sri- 
mangala, 2782 feet. South Coorg, peninsular India, collected Febru- 
ary 6, 1913, by G. C. Shortridge. 

Material examined. — "Cotengody Estate," 3500 feet [Nelliam- 
pathi Hills], Cochin (BM), four; "Shernelly," 1500 feet. Cochin 
(BM), four; Mudumalai, 3285 feet, Nilgiri Hills (AMNH), one; 
"Dirajpet," 3000 feet. South Coorg (BM), three; "Makut," 250 feet. 
South Coorg (BM), two; "Srimangala," 2782 feet. South Coorg (BM), 
one; "Wotekotti," 2000 feet. South Coorg (BM), one, (CNHM), two; 
"Benhope," 3000-4000 feet, Nilgiri Hills (CNHM), two. 

Original description. — ". . . clearly an ally of [tristriatus] . . . gen- 
eral colour greyish brown . . . three pale yellow longitudinal stripes 
. . . the middle one being much shorter and narrower than the lateral 
ones. . . . Saddle rich chestnut, this being one of the most striking 
characteristics . . . distinguished by its large size, the conspicuous 
chestnut color of the dorsal fur, and the black and white appearance 
of its tail. 

"Examples from N. Kanara and Dharwar decidedly smaller, and 
have a different general colour and are nearer to the type of F. tri- 

"The series of 35 . . . from Coorg is very constant in colouring, 
only three . . . having any black on the back, whereas the type, four 
of the five from Travancore, and about half of the specimens from 
N. Kanara have the saddle black ..." 

The pale stripes extend from shoulders to rump; the median 
stripe is 2 mm. wide; lateral ones are 3 to 4 mm. wide. The one ex- 
ample in the collections of the American Museum of Natural History 
has three bands of black on the tail hairs. 


Funatnbulus tristriatus numarius Wroughton 

Funambulus tristriattis numaritis Wroughton, 1916, Jour. Bombay Nat. Hist. 

Soc, 24, p. 646. 
Funambulus thomasi Wroughton and Davidson, 1919, Jour. Bombay Nat. Hist. 

Soc, 26, p. 729. 

Types. — Funambulus t. numarius, BM No., adult male 
from Helwak, Western Ghats, Satara District, Bombay, India, taken 
December 7, 1914, by S. H. Prater; thomasi, BM No., 
Khandala, 2000 feet. Western Ghats, Bombay, India, an adult female 
taken April 11, 1918, by Philip Gosse. 

Material examined. — "Potoli," 1800 feet. North Kanara (BM), 
four; Gersappa [Gersoppa], sea level, Kanara (BM), two; Helwak, 
Satara District (BM), eight, (CNHM), two (topotypes); "Hulekal," 
1500 feet, near Sirsi, Kanara (BM), one; Khandala, 1800 feet (BM), 
two (topotypes); Khed, Ratnagiri District (BM), two; Sirsi, 1500 
feet, Kanara (BM), one; Thana (BM), two; "Kardibetta Forest," 
2500 feet, Shimoga District (BM), one; Sagar, 2500 feet, Shimoga 
District (BM), one; "Devikop," 2000 feet. South Mahratta (BM), 

Original description. — "A local race of tristriatus, slightly smaller. 
. . . General colour above a grizzle of black, and buff giving a general 
effect of yellowish 'drab' approaching 'isabella colour,' the saddle . . . 
darker . . . with three longitudinal pale buff lines, broader and better 
marked than in true tristriatus. Face coloured like the back with a 
yellow suffusion, cheeks buff. In the summer coat the . . . saddle 
becomes jet black, and the dorsal stripes tend to become white, while 
the . . . face becomes tawny and the cheeks more ochraceous. Below 
dull white except the anal region, which like the under side of the 
tail, is a bright 'cinnamon rufous'; above the tail hairs are black with 
white tips, arranged ... to indicate ... a barring . . . 

"Shortridge obtained 35 specimens in Dharwar and Kanara which 
are clearly intermediates between [numarius] and tristriatus. . . ." 

Funambulus layardi (Blyth) 

Definition. — This species is constituted by the largest and most 
colorful squirrels of the genus Funambulus on Ceylon, subspecies 
layardi and signatus, and in the jungles of Travancore by the sub- 
species dravidianus. See the species distribution in Figure 10. 

Diagnosis. — The ventral pelage is entirely orange chestnut to 
cinnamon or may have the thoracic part yellowish orange. 





8°N + 

Fig. 10. Distribution of the Sinhalese jungle squirrel, Funambulus layardi, 
plotted from Ceylon localities recorded by Phillips (1935). Subspecies: A, layardi; 
B, signatus; C, dravidianus. 

Relationships to other species. — The species layardi seems to be 
the ecological equivalent on Ceylon of tristriatus in the Western 
Ghats. This is to say that species layardi is larger than forms of the 
species palmarum and sublineatus on Ceylon (see measurements given 
in pertinent subspecies accounts and in Table 3) and occupies prin- 
cipally the old evergreen rainforest. The fact that layardi seems 
also to occur in the southern Western Ghats, invites a comparative 
investigation of the ecology and distribution of these two species on 


the mainland. Hutton (1949, p. 692) reports the species layardi very 
common in the High Wavy Mountains of the southwest corner of 
Madura District, Madras, on the Travancore border, and tristriatus 
seems to be absent from this locality. 

Funambulus layardi layardi (Blyth) 

Sciurus layardi Blyth, 1849, Jour. Asiatic Soc, Bengal, 18, p. 602. 

Type.—IM No. 9481 (ASB 341 A), skin only, sex unknown, taken 
in the "uplands" of Ceylon in 1843 by E. L. Layard. 

Material examined. — "Mousa Kaida" [Mousakande, Gammad- 
uwa, Ceylon], 3000 feet (BM), one, (NMC), one; "Thimbulketiya," 
Ceylon (NMC), one. 

Type description. — The dorsal color of the type is now (1951) 
grizzled dark grayish brown. Of the three pale dorsal lines the 
median one is bright orange-yellow and 2-3 mm. wide; the lateral 
lines are only slightly paler than the body color and 3 mm. wide. 
The areas between the median and lateral lines are blackish brown. 
A faint dark line appears outside each lateral line. The median line 
extends from nape to rump and is 110 mm. long, the laterals from 
shoulder to rump, 85 mm. long. The head is colored like the back. 
The tail is grizzled like the body, its hairs with yellow-buff bases, 
long black subterminal rings, each 11 mm. in length, and buffy white 
tips. The end of the tail is black. The under parts are russet, the 
hairs with gray bases. 

Dimensions. — Phillips (1935, p. 237) offers averages and maxima 
of measurements taken in millimeters on three males and six females 
of F. I. layardi. The averages are respectively: length of head and 
body, 165 and 155.5; length of tail, 144 and 142; length of hind foot, 
37.5 and 37.5; length of ear, 16 and 15.5. The maxima are respec- 
tively: 168 and 160, 145 and 149, 39 and 39, and 16 and 16. 

Thomas (1924, p. 241) comments, ". . . the type of layardi came 
from the Ambigamoa Hills which are in the highlands of the Central 
Province (7°N.,80°30'E.) . . ." 

Funambulus layardi signatus Thomas 

Funambulus layardi signatus Thomas, 1924, Ann. Mag. Nat. Hist., (ser. 9), 
13, p. 241. 

Type. — BM No., young male from Rakwana, Ratna- 
pura District, 115 feet, Ceylon, collected February 17, 1914, by 
E. W. Mayor. 


Material examined. — Southeast of Ratnapura, Ceylon (BM), one. 

Original description. — "Similar in all respects to true layardi, but 
the light median dorsal line, instead of being narrow (about 3 mm.) 
and pale buffy, is broader (nearly 5 mm.) and of a rich ochraceous or 

Dimensions. — In Phillips (1935, p. 239) one finds averages of 
measurements taken in millimeters on eight males: length of head 
and body, 153.7; length of tail, 140.8; length of hind foot, 35.2; 
length of ear, 17. The maxima for the same series are respectively: 
160, 153, 38, and 19. 

Funambulus layardi dravidianus Robinson 

Funambulus layardi dravidianus Robinson, 1917, Rec. Indian Mus., 13, p. 42. 

Type. — IM No, 10143 (=9773), young individual from western 
side of the Western Ghats, Travancore, India, taken by Nelson 
Annandale. The reported presence of deciduous premolars in this 
type is the basis, acceptable to us, for regarding it as immature 
(e.g., Moore, 1956, pp. 46, 47). 

Material examined.— "India" (MNHN), one. 

Original description. — "Differs from [layardi] in having the top of 
the head and cheeks rich rufous orange, and under surface yellowish 
orange instead of dull chestnut. Area between light lines on back, 
deep lustrous black." 

Type description. — The general color of the upper parts are as in 
F. I. layardi of Ceylon, but the head is much more reddish and griz- 
zled with black. The dorsal area between and outside the three light 
lines is much blacker. The three lines are colored alike, light orange- 
buff. The width of the median line is 2 mm., of the lateral lines, 
3 mm. The hands, feet, and tail are as in layardi. The under parts 
differ sharply from the typical subspecies. The red of the crown be- 
comes almost clear chestnut on sides of the head and on either side 
of the throat. The throat and neck beneath are cinnamon, changing 
to yellow hairs (with gray bases) on the thorax. Behind this the 
pelage becomes dull brownish cinnamon (hairs with gray bases). 
The insides of the fore limbs are yellow-buff, of the hind limbs, 
brownish cinnamon. The under side of the base of the tail is brighter 
reddish cinnamon. 

Although we have been able to examine but two specimens of this 
form, Hutton (1949, p. 692) in reporting on it from the southwest 
corner of Madura District, Madras, on the Travancore border, re- 


garded it, "A very common little animal found throughout the hills 
above 3,000 feet." Among other comments he adds, "They often 
forage on the ground for insects." This was apparently in continu- 
ous evergreen jungle. 

Funambulus sublineatus (Waterhouse) 

Definition. — This species is constituted by the most diminutive 
forms of the genus Funambulus, and these are all endemic to the 
southern tip of peninsular India and to Ceylon as shown in Figure 11. 

Diagnosis. — (1) The fur is long and soft, tending to obscure the 
three rather faint light stripes. (2) The baculum is bifurcated and 
the tips end in rather spherical nobs. (3) The supra-orbital notches 
are obsolescent or absent. 

Relationships to other species. — On Ceylon Phillips (1935, p. 240) 
states clearly that sublineatus "... is confined to the jungles of the 
hill and wet zones and is not found at all in the dry zone." And 
(p. 241) he says, "Its favorite haunt is a dense bamboo brake or 
patch of . . . Strobilanthes sp. and undergrowth in the jungle. ... It 
spends almost as much time upon the ground as in the undergrowth, 
and though it [is observed] to examine the rough bark of large trees, 
it is but rarely seen in their upper branches." 

The larger species, layardi, occurs in these same wet jungles of 
the same central hill zone and southwestern zone of Ceylon as sub- 
lineatus, according to Phillips (1935, pp. 237-239), in ". . . damp, 
heavy forests of a medium altitude, from about 1,000 ft. to 4,000 ft.; 
it is found only in the tallest trees ... it seems to spend much of 
the day among the foliage of the tops of the tallest trees. ... It is, 
however, in no way averse to descending to the ground and may fre- 
quently be seen examining the boles of the forest giants or making 
its way through the undergrowth." For other differences between 
these two species in Ceylon, and comment on it, see "Intrageneric 
Relationships" in the above account of the genus Funambulus. 

Funambulus sublineatus sublineatus (Waterhouse) 

Sciurus sublineatus Waterhouse, 1838, Proc. Zool. Soc. London, 1838, p. 19. 

Sciurus delesserti Gervais, 1841, L'Institut, p. 171. 

Sciurits trilineatus Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 602. 

Types. — Sciurus sublineatus, BM No., young male 
from the Nilgiri Hills, Madras, India; delesserti, BM No. 217a, adult 
male from Nilgiri Hills, India; trilineatus, not found. 



Fig. 11. Geographic distribution of the dusky Indian jungle squirrel, Funam- 
bulus sublineatus, as indicated from material examined. Subspecies: A, subline- 
atus; B, obscurus. 

Material examined. — "Benhope," 3000-4000 feet, Nilgiri Hills 
(BM), one; "Bombay Shola," 7000 feet, Kodai Kanal [Kodaikanal], 
Palni Hills (BM), three; Coonoor, Nilgiri Hills (BM), two; "High 
Range," Trevandrum, Travancore (BM), one; "Madras" (BM), one; 
Manantoddy, Wynaad (BM), one; "Rookery Kil (Estate)," 3800 feet 
Kotagiri, Nilgiri Hills (BM), one; "Travancore," 4500-5000 feet 
(BM), two; "Huvinakadw Estate," 2843 feet, Kirtta, S. Coorg (BM), 
one; Kuttyani [Kuttyadi], 250 feet (BM), four; Ponmudi, 2000 feet 


(BM), two; "Shernelly," 1500 feet, Cochin, S. India (BM), one; 
"Tiger Shola," 5500 feet (BM), one; "Cumbum," 5000 feet, High 
Wavy Mts., Madura (CNHM), two; Ootacamund, Nilgiri Hills 
(UMMZ), one. 

Original description (translation from the Latin). — ^"Above fusco- 
olivaceous, washed with yellowish; four black dorsal lines, three whit- 
ish, running from shoulders to rump; under parts yellowish; tail 
ringed black and yellow. 

"... four dark and three pale lines on the back: these lines . . . 
are very narrow . . . are not continued onto the shoulders, [nor] over 
the haunches. The general colour olive-brown . . . throat, chest and 
rump are whitish, . . . belly is yellow ..." 

Pelage color. — The CNHM study skins are an agouti of about 
Dresden Brown and are uniformly so from snout to ankles excepting 
for the three light stripes which are only somewhat paler. The four 
dark lines embrace the light ones and are slightly wider. The tails 
seem not at all distichous and taper to a point. 

Habits. — Hutton (1949, p. 692) reporting on this form in the 
southwest corner of Madura District, Madras, on the Travancore 
border, remarks, "It does not like thick forests and so is not often 
seen in our jungles [on a plateau above 4000 feet]. It is quite com- 
mon in the Varushnaad [Valley], and in light evergreen forest. It 
prefers country that has a light rainfall and [is] fairly sheltered. 
In the forests at the south of the Varushnaad [Valley], however, it 
is quite common up to 5,000 ft." 

Funambulus sublineatus obscurus (Pelzeln and Kohl) 

Sciurus palmarum var. obscura Pelzeln and Kohl, 1886, Verhandl. Zool. Bot. 

Gesell. Wien, 35, p. 525. 
Funambulus kathleenae Thomas and Wroughton, 1915, Jour. Bombay Nat. 

Hist. Soc, 24, p. 38. 

Types. — Sciurus p. obscura, not seen, from the "Mountains of 
Ceylon, 1000 meters"; kathleenae, BM No., young male 
from Kottawa, Southern Province, Ceylon, collected April 11, 1913, 
by E. W. Mayor. 

Material examined, all from Ceylon.— Pattipola, 6210 feet. Cen- 
tral Province (BM), two; West Kaputale [Haputale], 6000 feet, 
Ohiya (NMC), one, (BM), three; Gammaduwa, 3400 feet, Mousa- 
kande (MCZ), one, (NMC), one. 

Original descriptions. — Sciurus p. obscurus, "From the mountains 
(1,000 meters elev.) ; the ones from the lowlands are of lighter color. 


The example under consideration is distinguished from the normal 
by the much darker coloring. The upper side is brown-gray; each 
hair shows a reddish yellow ring before its dark tip. The back with 
four dark chestnut-brown longitudinal stripes which are separated 
from one another by three much narrower pale yellow longitudinal 
stripes. The under side is lighter than the upper and shows more 

Funambulus kathleenae, "The readiest means of distinguishing 
kathleenae from sublineatus is the much greater width of the dark 
dorsal stripes, which are 7-8 mm. in breadth, as contrasted with 4-5." 

Habits. — This form is confined to Ceylon. According to Phillips 
(1935, p. 240), "... this little squirrel [occurs in] the jungles of the 
hills, from the highest peaks down to about 2,000 or 1,500 ft.; but 
... in the southwest generally, it descends to the lower foothills . . . 
through the damp, hilly jungle of the low-country ... to near the sea 
coast in [the Southern Province]. It is confined to the jungles of the 
hill and wet-zones and is not found at all in the dry -zone." 

Dimensions. — For dimensions of this form Phillips (1935, p. 240) 
presents averages and maxima of measurements taken in millimeters 
on eight males and eight females. The averages are respectively: 
length of head and body, 118 and 112.8; length of tail, 109.5 and 
103.5; length of hind foot, 28 and 30.3; length of ear, 14.4 and 14.5. 
The respective maxima are: 126 and 124, 117 and 113, 32 and 32, 
and 16 and 18. Maximum weight of either sex is 2}4 ounces. 

Genus CALLOSCIURUS Gray, 1867 

Callosciurus Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 277. 
Baginia Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 279. 
Erythrosciurus Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 285. 
Heterosciurus Trouessart, Le Naturaliste, 1 , p. 292. 
Tomeutes Thomas, 1915, Ann. Mag. Nat. Hist., (ser. 8), 15, p. 385. 

Type species. — Callosciurus, Sciurus rafflesi Vogors and Horsfield, 
the Sumatran form of C. prevosti Demarest from Malacca; Baginia, 
Sciurus notatus Boddaert from Java; Erythrosciurus, Sciurus ferru- 
gineus Cuvier from Burma; Heterosciurus, Sciurus erythraeus Pallas 
from Assam, India; Tomeutes, Sciurus lokroides Hodgson from Nepal 
a subspecies of C. pygerythrus Geoffroy from Burma. 

Definition. — Callosciurus is a genus of tree squirrels endemic to 
the Malaysian and Indochinese subregions, and is composed of the 
species erythraeus, ferrugineus, flavimanus, finlaysoni, caniceps, phay- 
rei, inornatus, and pygerythrus of the Indochinese Subregion (see 
their ranges in Figures 13-16) and extraterritorial species prevosti, 
notatus, nigrovitatus, albescens, and melanogaster of the Malaysian 

Diagnosis. — Callosciurus possesses the following characteristics: 
(1) The least interorbital breadth exceeds the greatest length of 
nasal. (2) There are no pronounced longitudinal stripes on the back 
(although there may be on the sides and venter). (3) There is a 
single, unforked, bony septum across the chamber of the auditory 
bulla. (4) The third upper premolar is present. (5) The coronoid 
process of the mandible is high and falcate. (6) The upper edge of 
the infra-orbital foramen is well separated from the maxillo-premaxil- 
lary suture. (7) The baculum consists of two separate parts, a shaft 
and a blade. (8) Orbit length exceeds 13 mm. (9) The supra-orbital 
notches are obsolescent. 

Callosciurus is distinguished from the other genera of the Indian 
and Indochinese subregions by the above characters as follows: from 
Ratufa by 3, 4, 6, and 7; Funamhulus by 2, 5, and 7; Tamiops by 



2 and 8; Dremomys 1; Menetes by 1 and 2; and Sciurotamias by 
1, 3, and 9. 

Intrageneric relationships. — Species of the genus Callosciurus are 
the common tree squirrels of the Indochinese and Malaysian sub- 
regions. No members of this genus cross the Garo-Rajmahal Gap 
into the Indian Subregion. Throughout a great deal of the Indo- 
Chinese Subregion there are two species of Callosciurus present in 
any one area. We recognize eight species of Callosciurus in the In- 
dochinese Subregion, and we consider them more closely related to 
each other than to any of the other species of Callosciurus, which 
occur only south of the Isthmus of Kra. The evidence so far ad- 
vanced (Moore, 1961a, p. 14) on which this monophyletally of the 
eight mainland species is inferred, is that the number of pairs of func- 
tional mammae is consistently only two; whereas the characteristic 
number of pairs in the several Malaysian species is three. This may 
be too weak a distinction to be regarded as subgeneric, but it does 
seem to distinguish what is a phylogenetic group higher than an ar- 
tenkreis and composed of two artenkreis. In further study this 
arrangement will probably be found to be supported in certain gen- 
eral characteristics of the pelage. For convenience until further 
study is accomplished, one of these two natural units may be called 
the mainland unit and the other the Sundaland unit of genus Callo- 
sciurus. There is some overlap in the distribution of these two units. 
The Malay Peninsula south of the Isthmus of Kra is occupied by 
two species of the mainland unit as well as three species of the Sunda- 
land unit. 

The mainland unit, we find, consists of one artenkreis of four allo- 
patric species spread over the whole Indochinese Subregion (eryth- 
raeus, ferrugineus, flavimanus, and finlaysoni) and a second artenkreis 
of four allopatric species {pygerythrus, phayrei, caniceps, and inor- 
natus) which is not so widespread but is on the whole sympatric with 
the first artenkreis. Although the species listed here bear names 
employed by Ellerman and Morrison-Scott (1951) and earlier auth- 
ors, the composition of our species usually differs markedly from 
earlier concepts. The reasons for the differences will be found in the 
following accounts. 

Descriptive skull characters. — (1) Least interorbital breadth ap- 
proximates orbitonasal length. (2) The occlusal plane is parallel to 
a plane connecting the tips of the upper incisors and the ventral 
extremities of the auditory bullae. (3) There is a subsquamosal 
foramen without vestige, or incipience, of a postglenoid foramen. 


Fig. 12. Skull and left mandible of the Irrawaddy squirrel, Callosciurus 
pygerythrus, AMNH No. 163507, X 1. Note the indication of a single septum 
dividing the auditory bulla. 

(4) The frontals are level along the line of least interorbital breadth, 
but rise to slight inflations where they meet the premaxillaries. (5) 
The temporal ridges do not form sagittal crests (excepting in a small 
percent of adult melanogaster) . (6) The temporal foramen is usually 
fairly well developed. (7) The jugal is broad with a fairly prominent 
postorbital process. (8) The lip of the infra-orbital foramen in lateral 
aspect is ordinarily concave. (9) The upper incisors are proodont. 

Callosciurus erythraeus (Pallas) 

Definition. — Callosciurus erythraeus is a species occupying the 
area of the Indochinese subregion west of the Irrawaddy River and 
its tributary the Nmai. It consists of the subspecies erythraeus, 
erythrogaster, bhutanensis, intermedius, sladeni, hartoni, and haring- 
toni and the named forms here included in these subspecies. See 
the species map in Figure 13. 

Diagnosis. — In this extraordinarily variable species no pelage (or 
other) characters are known which distinguish all of its subspecies 
from all the subspecies of its nearest relatives. See the accounts of 
the species flavimanus and ferrugineus for a discussion of the evi- 
dence of relationships between erythraeus as here recognized and its 
closest relatives. Table 4 provides some measurements of body and 
skull for 17 types of named forms of this species. 

Relationships to other species. — ^We have been at great pains with 
Callosciurus in the Indochinese Subregion to ascertain where there 
is evidence of intergradation between forms and where in absence of 
intergradation there is also positive evidence for recognizing alio- 



















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patric species. There is, of course, no question that the "Checklist 
of Palaearctic and Indian mammals" of Ellerman and Morrison- 
Scott (1951) is a work of enormous practical value to mammal ogists 
and that it will continue for many years to be as great a help to other 
as it has been to us in making this revision. But it is only a check- 
list, and its authors did not, and for such a scope could not, examine 
evidence in such detail as we have been able to do. Consequently, 
whereas they surely have made many shrewd taxonomic decisions, 
discovery that many other decisions are erroneous must be expected 
when detailed investigation of the available evidence is achieved. 
Such is the nature of the disagreement found here. The species of 
Callosciurus in the Indochinese Subregion that have been found in 
the present revision to be evidently good ones have in many instances 
the names used for species in the Palaearctic and Indian checklist. 
The subspecies which we find belong in these species, however, differ 
widely from those which Ellerman and Morrison-Scott (1951) in- 
clude, and this appears sometimes, as in the case of flavimanus, to 
result from rather basically different concepts. 

Species level difference between erythraeus and flavimanus is based 
on degree of difference in the areas of most probable contact, or most 
recent contact, and absence of evidence of intergradation. Dis- 
tinction at the species level from the other nearest related species, 
ferrugineus, is similarly based on degree of difference and absence of 
intergrades. Diagnostic difference between the two is that erythraeus 
has mostly agouti dorsal pelage, whereas ferrugineus has none and is 
all red. 

Throughout its entire range species C. erythraeus is (see Figure 13) 
sympatric with a smaller species C. pygerythrus (compare Figure 
16), but Morris (1936, p. 670) noted that there seemed to be some 
ecological or other factor along the Chindwin River enabling one or 
the other species to be the more abundant, ". . . from Singkaling 
Khamti southwards the east bank had been the productive area for 
[species erythraeus], the west bank producing mainly [species pyger- 
ythrus] which were rare on the east side. At Mawlaik the east bank 
again produced [erythraeus], only [pygerythrus] occurring on the west- 
ern side ..." 

Feeding habits of squirrel species in this subregion have been 
rarely noted, but Morris (1936, p. 671) recorded that, "At the time 
the expedition visited the [upper Chindwin River] area the squirrels 
[of these species] were observed to be feeding on Elaeocarpus sp., and 
Pterospermum sp." Although this statement is generalized, erythraeus 


is the commonest species encountered by the expedition and was evi- 
dently meant to be included; the implication is that they fed on the 
fruits of the trees mentioned. 

Callosciurus erythraeus erythraeus (Pallas) 

Sciurus erythraeus Pallas, 1778, Novae Species Quadrupedem Gliris Ordine . . ., 

p. 377. 
Callosciurus erythraeus wellsi Wroughton, 1921, Jour. Bombay Nat. Hist. Soc, 

27„ p. 775. 

Types. — Sciurus erythraeus, no longer in existence, the type local- 
ity restricted by Bonhote (1901a) to Assam, India, and here further 
restricted to the Garo Hills of Assam; wellsi, BM No., adult 
male, from Shangpung, 4000 feet, Jaintia Hills, Assam, India, taken 
July 10, 1920, by H. W. Wells. 

Material examined, all from Assam, India. — Tura (AMNH), 
three, (BM), three; Tura Mt., Garo Hills (CNHM), two; Umran 
(AMNH), five; Nongpoh, Khasia Hills (CNHM), one; Cherrapunji 
(CNHM), four, (UMMZ), five; Mawryngkueng, Khasia Hills (CN- 
HM), six; "Konshnong," 3000 feet, Jaintia Hills (BM), one, (CNHM) 
one; Duragiri, 3000 feet, Garo Hills (BM), one; "Mooston," 2000 
feet, Khasia Hills (BM), one; "Rajapara," 600 feet, south Kamrup 
(BM), five; "Kulsi," 150 feet, Kamrup (BM), one; "Shangpung," 
4000 feet, Jaintia Hills (BM), one. 

Pelade color. — The ventral pelage is generally Burnt Sienna (II), 
but in a few it is Chestnut and at least one Mars Orange. None has 
the ventral pelage divided, the throats are all red, and the chins re- 
main gray. The tails have the distal half or two-thirds red, some 
Burnt Sienna, some Mahogany Red. The ears are very bright. 
Ferruginous (XIV). The feet are very blackish agouti, contrasting 
rather strongly with the general dorsal pelage color (of back, sides, 
head, legs and proximal portion of the tail) an agouti of about Citrine 
Drab (XL). The tail has the same colors below as above. 

Discussion. — It appears that Blyth (1855, p. 473) identified this 
subspecies of the Garo Hills and Khasia Hills properly as the red- 
tailed erythraeus of Pallas, 1778, and at the same time distinguished 
the black-tailed subspecies to the east of it as a new species. These 
distinctions are perfectly good at the subspecies level in the mate- 
rial available to us. Wroughton's (1921) wellsi was described on a 
single character, a whitish tip to the end of the long red tip of the 
tail in the typical erythraeus. He reported this for only one locality, 
as constant in a series of nine specimens. As Blyth (1855, p. 473) 


had stated much earlier of typical erythraeus "... the terminal two- 
thirds or more of the tail are nearly of the same colour as the belly, 
the tip generally being paler." Some variation in the amount of this 
paler tipping is observed in the material, and constancy of excessively 
pale tips in a sample of nine from one locality, or even a cluster of 
localities, within the area of general variation is feeble characteriza- 
tion for a subspecies. 

CNHM No. 82847 taken at Syndai, 2500 feet elevation, in the 
Jaintia Hills is black-tailed like erythrogaster from the Lushai Hills, 
although material from other Jaintia Hills localities agrees in every 
way with C. e. erythraeus material from the Garo Hills and Khasia 

Callosciurus erythraeus erythrogaster (Blyth) 

Sciurus erythrogaster Blyth, 1842, Jour. Asiatic Soc. Bengal, 11, p. 970. 

Macroxus punctatissimus Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 283. 

Callosciurus erythraeus nagarum Thomas and Wroughton, 1916, Jour. Bom- 
bay Nat. Hist. Soc, 24, p. 228. 

Callosciurus erythraeus kinneari Thomas and Wroughton, 1916, Jour. Bom- 
bay Nat. Hist. Soc, 24, p. 229. 

Callosciurus erythraeus crotalius Thomas and Wroughton, 1916, Jour. Bombay 
Nat. Hist. Soc, 24, p. 229. 

Types. — Sciurus erythrogaster, IM No. 9262, collected in "Muni- 
pore" [Manipur], India, by C. S. Gullevie; punctatissimus, BM No., juvenile, from "India"; nagarum, BM No., 
adult male, from near Sadiya, northeast Assam, India, collected in 
April 1877, by A. 0. Hume; kinneari, BM No., an old male 
taken at Tatkon, near Kindat, Burma, west of the Chindwin River 
on June 26, 1914, by G. C. Shortridge; crotalius, BM No., 
an adult male from Hkamti, 500 feet, on the Chindwin River, in 
Burma, August 7, 1914, by G. C. Shortridge. 

Material examined, from Assam. — Aijal, Lushai Hills (CNHM), 
two; Hnathial, Lushai Hills (CNHM), one; "Syndai," Jaintia Hills 
(CNHM), one, (BM), three; Kohima, Naga Hills (CNHM), one, 
(UMMZ), one; Karong, Manipur (CNHM), one, (UMMZ), one; 
"Aimole," Manipur (BM), four; "Machi," Manipur (BM), two; 
"Noong-Zai-Dau," Manipur (BM), one; "Koomberong," Manipur 
(BM), one; "Dilkoosha," Cachar (BM), two; "Cachar" (USNM) 
one; Margherita, 400 feet, Naga Hills (CNHM), one; 13 miles east 
of Ledo (USNM), one; Dibrugahr, Rungagora (MCZ), one, (CNHM), 
one; Mokokchung, 4500-5000 feet (BM), two; Mikir Hills (BM), 


one; Tekhubama (BM), two; "Cholimsen," 4000 ft. (BM), one; 
"Langting," 1500 feet, Cachar Hills (BM), one; Takubama, Naga 
Hills (CNHM), one; Powai [Poi], 400 feet (BM), one. 

Material examined, all from west of the Chindwin River, Burma. — 
Mt. Victoria, 1400, 2200, and 2500 meters (AMNH), 16; opposite 
Homalin (AMNH), one; Tempao (AMNH), one; Pebin (AMNH), 
one; opposite Moklok (AMNH), one; Nauswa (AMNH), three; op- 
posite Kaungheim (AMNH), 19; opposite Limpa (AMNH), two; 
opposite Heinsum (AMNH), five; Tanai Hka, Dagung Hka (AMNH), 
one; Hahti (AMNH), two; "Haingyan," Naga Hills (BM), one; 
Tatkon, west of Kindat, Chindwin R. (AMNH), one, (CNHM), 
two, (BM), six; 50 to 65 miles west of Kindat, at 4000 to 5000 feet 
(BM), three; Tamanthe, 430 feet, upper Chindwin (BM), three; 
opposite Hkamti, Chindwin R. (BM), eight, (AMNH), one; Hai 
Bum, Chindwin R. (AMNH), 27; Chenga Hka, Chindwin R. 
(AMNH), 13; Dagung Hka, Chindwin R. (AMNH), two; Lachu 
Ga, Chindwin R. (AMNH), one; Taga Hka, Chindwin R. (AMNH), 
one; Lahkaw Hka, Chindwin R. (AMNH), one; 40 miles northwest 
of Kindat at 600 feet (BM), one; "Nanthalet," Kawbaw Valley 
(BM), one; Shingbwiyang, Hukawng Valley (BM), one; 20 miles 
north of Hkamti (BM), one. 

Type description. — A color description of the type of erythrogaster 
Blyth, 1842, recorded in 1959 follows: Ventral pelage of body and 
limbs entirely Mahogany Red excepting for chin which is invaded 
by agouti. Dorsal pelage a very punctate agouti of about Olive- 
Brown (XL). The guard hairs of the dorsum generally have two 
light bands per hair. The bands are a pale buff and one mm. or less 
in length. The more distal band is generally subapical and 3 to 5 mm. 
from the proximal light band, and the remainder of the hair is black. 
The variation in dorsal pelage color is very slight, the feet being 
blacker but still agouti, and the head very slightly browner. The 
ears appear to have been faintly redder than the head. The tail is 
agouti for one-third its length but blackens distally because the black 
tips of the hairs increase in proportional length. The tail hairs have 
as many as five light bands on a hair, each about two mm. long and 
separated about 5 mm. rather evenly. The head gradually lightens 
anteriorly until the snout and lips are Tawny Olive (XXIX). 

Pelage coZor.— The agouti dorsal pelage of back, sides, legs, tail, 
and head is about Grayish Olive (XLVI) in material from Manipur 
and the Naga Hills. The feet are slightly darker. The tail is black 
distally from one-third to all of its length, and in the hills the length 


of black seems clinal, least black in the north and increasing south- 
ward. The ears are brightest in the Manipur material, Hazel (XIV), 
but in the Lushai Hills material are about Chestnut Brown. Ven- 
tral pelage of body, throat, and limbs is generally all red, but varies 
slightly: Chestnut (II) in Takubama specimen. Auburn in Kohima 
one. Mahogany Red from Karong. The series of 16 from Mt. Vic- 
toria vary between Mahogany Red and Burnt Sienna. The Taku- 
bama specimen, which is from about the middle of the subspecies 
range, has an agouti throat, and the red ventral pelage is bilaterally 
divided full length by a 3 mm. stripe of agouti which is as gray as 
its back. The divided venter seems to occur only occasionally in 
this subspecies. In the (AMNH) series of 81 specimens from the 
many localities along the west bank of the Chindwin River only 12 
show even a definite incipience of this condition. Proximally the 
ventral side of tail is like the dorsum in color. The number of black 
bands on tail hairs seems to range from three (Mt. Victoria) to five 
(Lushai Hills). 

Discussion — In this treatment of erythrogaster several potential 
subspecies bearing other names are synonymized with it because 
they have the distinctive characters of erythrogaster. Although they 
also appear to have small divergent characters of their own, their 
known geographic range is very limited, and it seems definitely pref- 
erable to emphasize their relationship to erythrogaster by lumping 
them with it than to exaggerate their distinctiveness by giving them 
the same taxonomic rank. 

We have seen one (USNM) specimen from Cachar representing 
the named form punctatissimus. Its punctate dorsal pelage is about 
Bone Brown (XL), its ventral pelage Claret Brown (I). Feet punc- 
tate black; digits solid black. Tail black; tail hairs have four light 
bands obscured by long, black tips. The throat is punctate, and this 
extends posteriorly as a narrow wedge in the midline. Two speci- 
mens from Aijal, Lushai Hills, are similar enough to this to suggest 
the possibility of a blacker subspecies than erythrogaster occupying 
the monsoon rainforest of the area of west-facing slopes southward 
from Cachar possibly even along the Arakan Yoma. The large 
series of ordinary erythrogaster from Mt. Victoria appears to miti- 
gate against the existence of such a black subspecies, and there is 
a good possibility that these very black punctate specimens repre- 
sent strongly melanistic individual variants in a population of ordi- 
nary erythrogaster. The problem merits wider collecting. 

The named form kinneari is known only from Tatkon, and from 
Ahlaw and Nanthalet in the Kawbaw Valley probably 30 miles to 


the west, while from 50 to 65 miles west of Tatkon only ordinary 
erythrogaster (three specimens) are available. Furthermore, the char- 
acter supposed to distinguish kinneari occurs sporadically as an in- 
dividual variant farther north along the west bank of the Chindwin 
River: four among 26 specimens from Haibum, four among 19 from 
opposite Kaunghein, two taken between Jantang and Dagung Hka, 
and one each at Taga Hka and Lahkaw Hka. Reference to the ma- 
terial listed here and to Carter's (1943, frontispiece) map of these 
localities will show that more ordinary erythrogaster were collected 
at other localities between the above. This extremely odd kinneari 
character is surely worthy of wider collection and further study, 
but as now known, it does not appear to characterize a substantially 
geographic subspecies. 

The claim to subspecies status of crotalius rests on the distinc- 
tion of an inconspicuous white tip in the black distal portion of the 
tail. This is reported to be constant in 29 specimens from the type 
locality, Hkamti, by the describers of crotalius, and Carter (1943, 
pp. 103 and 110) indicates that it continues constant in occurrence 
in a sample of 48 more from Hkamti northeastward through seven 
other localities along the west side of the Chindwin River, a straight- 
line distance of 42 miles on Carter's (1943, frontispiece) map. See 
the paragraph beginning this discussion. 

The type specimen of nagarum, which is from Sadiya, does have 
a black tail although from (Tate's) color notes it seems otherwise 
very close to intermedins and would, of course, be intermediate be- 
tween the black-tailed race and intermedius. It seems likely that 
the type of nagarum may have come from opposite Sadiya across the 
Luhit River which for part of its length appears to be a barrier sep- 
arating the black-tailed subspecies erythrogaster on the south from 
intermedius on the north. The type of aquilo, also recorded as from 
Sadiya, was evidently taken beside the Dibong River north of Sadiya, 
and it agrees in critical pelage characteristics with the type of in- 

Callosciurus erythraeus bhutanensis (Bonhote) 

Sciurus erythraeus bhutanensis Bonhote, 1901, Ann. Mag. Nat. Hist. (ser. 7), 

7, p. 161. 
Callosciurus crumpi Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, 

p. 425. 

Types. — Sciurus e. bhutanensis, skull BM No., skin BM 
No. 41.1216, "East India Company" collection from "Bhutan"; 


crumpi, BM No., an adult male from Sedonchen, 6500 
feet, Sikkim, India, taken on November 17, 1914 by C. A. Crump. 

Material examined. — Lingtam, Sikkim (CNHM), four; Sedon- 
chen, Sikkim (BM), three. 

Pelage color. — The ventral pelage is all agouti, but there is a red 
suffusion in it which demarks the midventral division common in 
some other forms of erythraeus. The rostrum is all Cinnamon 
Rufous (XIV). The tail has a black tip. The feet are blackish 
agouti. The ears have a little orange color. The dorsal pelage is 
between Deep Olive (XL) and Olive Brown. The tail hairs have 
black tips and four black bands. 

Discussion. — It now appears from the type descriptions of hhu- 
tanensis and crumpi that the type of bhutanensis may be an inter- 
grade between erythrogaster and "crumpi." It is unfortunate (but 
not rare) to have the type locality in an area of intergradation. In 
this instance the type specimen has the red face which characterizes 
its race but has a venter like that of erythrogaster. 

Robinson (1913, p. 89) recorded two specimens from Pasighat on 
the west bank of the Dihang at about 28° 05' north latitude as of 
this subspecies. One of two other specimens taken about 12 miles 
west of Pasighat in the Abor Hills he identified as erythrogaster, the 
other he thought intermediate between erythrogaster and intermedius. 
There is an island in the Dihang River at Pasighat and much braid- 
ing immediately downstream which might well have functioned to 
disconnect pieces of land inhabited by intermedius from the east side 
of the main stream and eventually connect them with the west side, 
thus accounting for the presence of the above-mentioned intermedi- 
ate. Robinson (1913, p. 89) also reported a series of seven inter- 
medius from Kobo, which is on the plains just across the island-laden 
Dihang from Sadiya, which may represent a population that has 
reached the west side in this same fashion (or may actually have 
been collected from a population on an island near Kobo). 

The presence of erythrogaster in the Abor Hills north of the Brah- 
maputra River seems a piece of zoological evidence which supports 
the theory that the Dihang was formerly tributary to the Chindwin 
River (see Chhibber, 1934, p. 33) and that there was no great river 
barrier between the Abor Hills and the Naga Hills range of erythro- 
gaster until the Brahmaputra extended its headwaters and diverted 
the Dihang to its own channel. 

Habits. — Crump's field notes and speculations are worth quot- 
ing: "Found in heavy forest, singly or in pairs. When alarmed 


it usually makes for ground, it was also observed to come down in 
the evening, so probably it breeds in holes among the roots of trees. 
The call is a deep-toned clack rapidly repeated. I saw this squirrel 
only at Sedonchen." (C. A. Crump in Wroughton, 1916a, p. 487). 

Callosciurus erythraeus intermedius (Anderson) 

Sciurus gordoni, var. intermedia Anderson, 1879, Zool. Anat. Res. Yunnan, 

p. 241. 
Callosciurus castaneoventris aquilo Wroughton, 1921, Jour. Bombay Nat. Hist. 

Soc, 27, p. 601. 

Types. — Sciurus gordoni intermedia, IM No. 9260, a male from 
Dikrang Valley, Assam, India, taken by H. H. Godwin-Austin (type 
selected by H. C. Robinson, 1918, p. 198 footnote); aquilo, BM No., adult female, from Dibong River, 500 feet, Sadiya, north- 
west Assam, India, collected November 30, 1919, by H. W. Wells. 

Material examined, from Assam. — Sadiya (AMNH), two; Dreyi, 
5140 feet (AMNH), two, (BM), two; Tezu, 648 feet (AMNH), two; 
Dening, 2250 feet, Mishmi Hills (CNHM), two, (BM), nine; Raja- 
para, 600 feet, south Kamrup (BM), eight; "Lonit Valley," 800 feet, 
Sadiya (BM), one; "Tiki," 1440 feet, Mishmi Hills (BM), one; 
"Ain," 1500 feet, Mishmi Hills (BM), one. 

Material examined, from upper Burma. — Nam Tamai, 3000 and 
4000 feet, N. Myitkyina Province (USNM), two, (BM), eight; Nam 
Tamai Valley, 3500 and 6000 feet (CNHM), three; Adung Valley, 
5000 feet (CNHM), one; Hkamti Plain, 1500 feet, Kachin Province 
(CNHM), two; 10 miles E. of Hkamti Plain, 4000 feet (CNHM), 
one; Sumpra Bum, 2500 feet (CNHM), one; Kadrangyong, 2000 
feet, 75 miles north of Myitkyina (BM), one; Htinenan [Htingwan], 
3200 feet (BM), one; Lunghkang, 3000 feet (BM), two; Matsa- 
tap, 3500 feet (BM), one; Yunghtang, 4500 feet (BM), one; Sumka 
Uma, 2000 feet (BM), one; Taron Valley, 4000 feet (BM), one; 
Ratnamhti, 2500 feet (BM), one. 

Type description. — A description made of the type in 1959 fol- 
lows: Ventral pelage Mahogany Red except for throat which is agouti 
but partially infiltrated by the reddish color. The reddish agouti ex- 
tends as a midventral wedge the length of the thorax. The throat, 
anteriorly where not infused with red, is Citrine Drab (XL). The 
muzzle is about Isabella Color (XXX). The tail seems about the 
same color ventrally as dorsally, grossly Cinnamon-Brown. Proxi- 
mally the dorsum of the tail is, like the middorsum, just about 
Russet. The middorsum color extends to the rostrum but the red 


decreases anteriorly. Dorsal pelage of sides and legs Buffy Brown 
(XL). Digits all blackish agouti, about Mummy Brown (XV). 
Dorsal guard hairs have two reddish buff light bands each fully 
two mm. long and separated by a three mm. black band. Tips 
black. Tail hairs have reddish buff tips and four blackish bands. 
Ears and obscure eye-ring about color of muzzle. Tail not percep- 
tably distichous below. 

Diagnosis. — The subspecies intermedins differs from michianus to 
the east principally in having the middorsal agouti pelage infused 
with reddish instead of black. Further, the ventral aspect of the 
tail of michianus is notably lighter than that of intermedins. The 
subspecies intermedins differs from gordoni by lacking the incipient 
black tip of tail which characterizes gordoni, by usually lacking the 
agouti throat and even a partial agouti stripe like that which en- 
tirely bisects the red venter longitudinally in gordoni, and by ordi- 
narily possessing red ventral pelage extending to the very heel of the 
manus rather than red pelage separated from the manus by a centi- 
meter or two of agouti as in gordoni. The differences between inter- 
medins and erythrogaster and bhntanensis have been stated in the 
treatment of those subspecies. 

Discnssion. — The characters of the material representing Callo- 
scinrns sladeni vernayi (Carter, 1942) , appear to represent intergrada- 
tion between what has been lately known as Calloscinrus sladeni 
rnhex and Calloscinrns erythraens intermedins. The evidence is this: 
The tail in subspecies sladeni {= rnhex) has a long red tip very much 
as in subspecies erythraens, but between the ranges of intermedins 
and sladeni from N'bunghku to Dalu the Vernay-Hopwood expedi- 
tion collected this "vernayi" material with a plain, agouti tail like 
that of intermedins. The pes and manus are not as entirely red as 
in sladeni, but are invaded by agouti in varying amounts suggesting 
influence of the all agouti foot of intermedins. The amount of red- 
dish in the pelage of the dorsum is intermediate between that of in- 
termedins and sladeni. The scarcity of occurrence in intermedins of 
a midventral agouti stripe bisecting the red ventral pelage (despite 
the geographic location of this subspecies between other subspecies 
characterized by such a stripe — bhntanensis to the west and michi- 
anus and gordoni to the east) suggests that intermedins receives by 
interbreeding with sladeni the hereditary factor(s) for undivided 
venter which prevail (s) throughout the forms previously regarded 
as constituting the species sladeni. 


The color of the dark, black-tailed erythrogaster on the west side 
of the Chindwin River contrasts strikingly with the light or red- 
tailed subspecies on the east side. This extreme degree of difference 
indicates that these squirrels on the two sides of the Chindwin River 
have been effectively prevented from mingling and interbreeding for 
a long time. Where the Chindwin is a large and substantial river, 
and to an unknown extent crocodiles may augment its effectiveness 
as a barrier to passage of tree squirrels, the river barrier seems a 
good enough explanation of the observed differences. Up where the 
river narrows near its headwaters, however, the striking difference 
continues. At Dalu, the point farthest up-river (north) from which 
the Vernay-Hopwood expedition obtained squirrels of this species 
on both sides, sladeni shows definite intergradation with the grayer 
northern subspecies intermedins. But sladeni shows not the slightest 
evidence of intergradation with the strikingly different erythrogaster 
across, or around the head of, the river. 

In "The Geology of Burma" Chhibber (1934, p. 33) reports and 
illustrates a geologically earlier condition in which the Tsangpo River 
of Tibet flowed eastward into the Dihang River of Assam, which 
flowed southward into the Chindwin River of Burma. The principal 
physiographic changes which need to have taken place subsequently 
to reach present conditions are: 1. the head erosion by the Brahma- 
putra River of Assam, India, to capture the Dihang and its tributary 
Tsangpo; 2. sufficient uplift to account for a divide with the least 
elevation of 900 meters presently between the Dihang-Brahmaputra 
and the Chindwin drainages. The supposed enormously greater 
former length and volume of the Chindwin River would have pro- 
vided so impassable a physiographic barrier to spread by tree squir- 
rels, as to permit the observed differentiation of the species erythraens 
on the east and west sides of the river in Burma. The known dis- 
tribution of erythrogaster remains nicely west of the supposed former 
connection between the Dihang and the Chindwin, and shows no 
evidence of intergradation with intermedius even though the sup- 
posed former barrier is not there now. This may be evidence for 
recognizing them as separate species, but no specimens of interme- 
dius from closer to the supposed former barrier than 60 or 70 miles 
have been examined, and a narrow zone of intergradation may exist. 
Suggestion that intergradation probably does occur here is seen 
in the degree of difference in pelage color characteristics which is 
greater between intermedius and sladeni (which are shown above to be 
connected by intergrades) than between intermedium and erythrogaster. 


The observed geographic differentiation in the squirrel species 
Callosciurus erythraeus thus supports the hypothesis that the Tsang- 
po-Dihang formerly was part of the Chindwin River. It would be 
interesting in this connection to know to what extent the population 
of the species C. erythraeus which certainly must exist in Assam north 
of the Brahmaputra River (in the Dafla Hills, the Miri Hills, and 
especially the Abor Hills) has differentiated from the population 
south of that river (erythrogaster) since they were effectively sepa- 
rated by this stream piracy. 

Habits. — Field observations of subspecies intermedius are con- 
tributed in a letter of June 10, 1961, by Lord Cranbrook from his 
field notes on the Adung Valley expedition. "No. 63. Shot about 
20 feet up in low jungle. No. 56 low down in bushy tree. Nos. 74 
and 75. I watched them for a considerable time chasing each other 
about in the same few trees in fairly heavy jungle. Three times in a 
period of about % hour they came right down onto the ground, com- 
ing down slowly bit by bit and when on the ground constantly alert, 
running towards or even a foot or two up a tree at the slightest hint 
of danger, even the chattering of a babbler. These squirrels are very 
common. By sitting quite still almost anywhere in the jungle one 
can nearly always see one within half an hour at the longest. If 
alarmed they freeze for minutes at a time, often in what appear to 
be the most uncomfortable positions. Not bad eating. Dec. 30, 
1930. Crossing the ridge between the Thi Hka and Nam Tisang 
[see Cranbrook's itinerary in Kinnear (1934)] C erythraeus seen at 
5000 feet. Jan. 7, 1931. Nam Hat-Nam Tamai divide. Several 
C erythraeus seen at 4 to 5000 feet and two collected, so they must 
be common. The valleys of the Nam Tisang and Nam Hat are very 
damp; full of leeches and blister flies which may account for not hav- 
ing seen any squirrels in them. The Hkampti Plain where they were 
very common is comparatively dry at this time of year. No. 147 
pregnant January. Common in Nam Tamai valley." 

Callosciurus erythraeus sladeni (Anderson) 

Sciurus sladeni Anderson, 1871, Proc. Zool. Soc. London, 1871, p. 139, pi. 20. 
Sciuriis kemmisi Wroughton, 1908, Ann. Mag. Nat. Hist. (ser. 8), 2, p. 491. 
Sciurtis sladeni rubex Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, p. 198. 
Sciurus sladeni midas Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, p. 198. 
Callosciurus sladeni vernayi Carter, 1942, Amer. Mus. Novitates, no. 1208, p. 1. 

Types. — Sciurus sladeni (not seen) IM No. 9371, a male taken 
at Thizyain [Tigyaing], Burma, on the upper Irrawaddy River Jan- 


uary 18, 1868, by John Anderson (according to Robinson and Kloss, 
1918, p. 201) ; kemmisi, BM No., adult female (skull missing), 
from Katha, upper Irrawaddy, Burma, collected by A. W. Kemmis; 
ruhex, BM No. 14.4.37, adult male, from Lonkin, Myitkyina Dis- 
trict, north Burma, collected February 22, 1914 by F. E. W. Venning 
(Ellerman and Morrison-Scott, 1951, p. 482, proposed to change this 
to Yin on the Lower Chindwin River, asserting that the animal does 
not occur at Lonkin. Yet the Vernay-Hopwood Expedition brought 
back two, AMNH Nos. 113235 and 113241, from Lonkin, collected 
in January. The face in one is clear orange red as in Thomas' type, 
that of the other much more grizzled); midas, BM No., 
old male, from Myitkyina District, 600 feet, north Burma, collected 
May 2, 1911 by A. W. Kemmis; vernayi, AMNH No. 113252, adult 
female, from Tapa Hka, northwest of Myitkyina, north Burma, col- 
lected February 2, 1935, by Vernay-Hopwood Chindwin Expedition. 

t Material examined, all from east of the Chindwin River, Burma. — 
Yin (BM), eight; opposite Kalewa (AMNH), six; Taukchandmai, 
500 feet, 20 miles SE of Kindat (BM), one; opposite Mawlaik 
(AMNH), four; Kindat, 250 feet (BM), 11; Pantha (AMNH), one; 
Wuntho (BM), one; Katha, 1000 feet (BM), one; "Changzon," 1000 
feet, Shweli River (BM), one; "Nyoung Bintha," left bank of Irra- 
waddy River (BM), one; "Myitkyina District," 600 ft. (BM), one; 
"Kadranyang," Fort Hertz Road (BM), one; Nanyaseik (AMNH), 
10; Lonkin (AMNH), four; Tawmaw (AMNH), four; Mansum 
(AMNH), one; N'bunghku (AMNH), two; Pumsin (AMNH), four; 
Zaulep Ga (AMNH), one; Tapa Hka (AMNH), four; Dalu (AMNH), 

Discussion. — The medium-sized tree squirrels occupying the phys- 
iographically varied area in upper Burma which lies between the 
Chindwin and Irrawaddy rivers, show extreme variation in pelage 
characters which is unquestionably geographic. They also possess 
two pelage characters which show them to be, so far as noticed, more 
closely related to each other than to medium-sized squirrels outside 
of this area. The pelage characters linking these squirrels together 
are: rostrum and forward part of crown colored like feet; dorsal color 
of manus continuing onto the forearm for half the forearm length. 
These characters, we presume, have led some earlier students, not 
unjustifiably, to consider these squirrels a single species, Callosciurus 
sladeni, and to regard, often with equally good justification, its geo- 
graphical variants as subspecies. In the account of C. erythraeus 
intermedius, however, we have reported evidence that sladeni inter- 


grades with intermedius, and we therefore recognize the old species 
sladeni as a series of subspecies of erythraeus. 

Between subspecies sladeni and haringtoni a related complex of 
squirrels occupies a curved, band-like area hemming the nearly white 
subspecies haringtoni against the Chindwin River. Within this band 
there is some justification for the naming of material from the south 
millardi as a subspecies distinct from the northern shortridgei, but 
bartoni is intermediate between them in color of pelage as well as 
geographically. In the American Museum material, no very real 
justification is seen for the distinction of careyi from fryanus or fry- 
anus from shortridgei. Furthermore, it would unnecessarily confuse 
the more important relationships to accept fragmentation of the 
squirrels of this band-like area into two or more subspecies, even on 
a north-south basis, unless it could be demonstrated that the Uyu 
River or some other partial barrier causes a fairly sharp break in the 
intergradation of their pelage characters. The important relation- 
ships of this band are that it intergrades with haringtoni on the west 
and with sladeni on the northeast and south. Every pelage character 
of the bartoni complex is either intermediate between haringtoni and 
sladeni or is also possessed by either haringtoni or sladeni. Table 5 
shows some indication of this intergradation. 

Table 5 lists the localities at which the Vernay-Hopwood Expe- 
dition collected specimens of the squirrel that has earlier been re- 
garded as the species sladeni. These localities are distributed in the 
table in a manner roughly approximating their geographic distribu- 
tion, the top of the table representing north, the bottom south. The 
eastern localities in the upper part of the column, however, are ac- 
tually distributed along a line of march which is grossly northwest 
from Nanyaseik to Tapa Hka, and the other series extends in a line 
almost southwest from Dalu to Kalewa (Carter, 1943, map frontis- 
piece). Because the eastern localities are all somewhat equidistant 
from the more southern Chindwin localities, their relative northern- 
ness may be taken as approximating that of Dalu for purposes of 
the table. 

The distances given in Table 5 are approximate airline miles 
(taken from Carter's 1943 frontispiece map) for the distance be- 
tween each locality and the one above it. Where two figures are 
given separated by a comma in the table, both numbers of bands 
were found, and the first number predominated. Ten to twenty 
hairs were plucked from the mid-sagittal parts of the body pelage 
(indicated in Table 5) of each specimen and their color bands counted 


Table 5. Variation in number of color bands on hairs of named forms at 
collecting stations of Vernay-Hopwood expedition. (Data are for black 
bands on tail hairs, light bands on other. Detailed explanation in text.) 




named forms 





Tapa Hka 
























































S. Hkamti 






















































































under magnification. Bands on tail hairs were counted in situ about 
at the midlength of the tail. Each locality is represented by one 
specimen, selected from the available material for having tail hairs 
fully grown out and dorsal pelage fresh and unworn. 

With the aid of the above explanation of Table 5, indications 
may be seen that the number of bands in the hairs decreases from 
the sladeni {=vernayi=ruhex) material in the north through the 
northern extremity of the bartoni (=shortridgei=fryanus= careyi) 
band to haringtoni. Continuing southward from haringtoni through 
the southern extremity of the bartoni {=millardi) band, which is 
regrettably not represented in the Vernay-Hopwood Expedition col- 
lection, to sladeni again one finds the number increased. 

By this lumping, which may seem crude in view of the striking 
characters involved, one emerges with an understandable, if some- 
what hypothetical, distributional variation in the forms of what has 


been regarded as the species sladeni. For an airline distance of 22 
miles along the Chindwin River the astonishingly light, cream-col- 
ored form haringtoni is known to occur. Although the Chindwin 
River axis of its range may eventually prove to be somewhat greater 
than 22 airline miles, the latitudinal axis of its range does not extend 
directly east more than 22 miles beyond the Uyu River. This range 
is extremely small for a form accepted here as a good geographic sub- 
species, but this is one of the most strikingly distinct subspecies in 
the whole retinue of the greatly variable species erythraeus. Sur- 
rounding haringtoni, together with the Chindwin River, is the sub- 
species bartoni which forms a band probably no more than 40 miles 
wide and 150 long. The squirrels of this band are variable but cer- 
tainly not variable enough geographically to justify their fragmen- 
tation into the five named forms. Outside of the band which is 
bartoni lies the subspecies sladeni. This occupies the suitable habitat 
of the remaining area between the Chindwin and Irrawaddy rivers. 
Subspecies sladeni is variable also, but the variation observed in it 
that is geographic does not justify retention of the five named forms. 

Callosciurus erythraeus bartoni (Thomas) 

Sciurus sladeni bartoni Thomas, 1914, Jour. Bombay Nat. Hist, Soc, 23, 

p. 198. 
Callosciurus sladeni shoriridgei Thomas and Wroughton, 1916, Jour. Bombay 

Nat. Hist. Soc, 24, p. 232. 
Callosciurus sladeni fryanus Thomas and Wroughton, 1916, Jour. Bombay 

Nat. Hist. Soc, 24, p. 232. 
Callosciurus sladeni millardi Thomas and Wroughton, 1916, Jour. Bombay 

Nat. Hist. Soc, 24, p. 233. 
Callosciurus sladeni careyi Thomas and Wroughton, 1916, Jour. Bombay Nat. 

Hist. Soc, 24, p. 233. 

Types. — Sciurus s. bartoni, BM No., old female, from 
Uyu River, 900 feet, 20 miles northwest of Mansi, upper Burma, col- 
lected March 24, 1911 by C. S. Barton; shortridgei, BM No. 15.5.5.- 
104, adult female, from Hkamti, Upper Chindwin River, north 
Burma, collected July 28, 1914 by G. C. Shortridge and S. A. Mac- 
millan; fryanus, BM No., young female, from Munsin, 
450 feet, upper Chindwin River, north Burma, collected August 14, 
1914, by G. C. Shortridge and S. A. Macmillan; millardi, BM No., old male, from Paungbyin between Kindat and Homalin, 
upper Chindwin River, north Burma, collected June 21, 1914 by 
G. C. Shortridge and S. A. Macmillan; careyi, BM No., 
old female, from Tamanthe, 430 feet, upper Chindwin River, north 


Burma, collected August 16, 1914, by G. C. Shortridge and S. A. 

Material examined, all from east of the Chindwin River, Burma. — 
Pyaungbyin (BM), two; Intha, Chindwin River (BM), two; Man the 
(AMNH), one; Tamanthe (AMNH), one, (BM), four; Phawzaw 
(AMNH), four; Minsin, 450 feet (BM), four; Moklok (AMNH), 
two; Kaunghein (AMNH), 26; Kauktaung, 475 feet (BM), two; 
Limpa (AMNH), three; Heinsum (AMNH), four; Singkahng Hkamti 
(AMNH), 40, (BM), 11, (CNHM), two. 

Diagnosis. — The characteristics distinguishing this admittedly 
heterogeneous subspecies from its geographic neighbors are: (1) The 
dorsal pelage of rostrum and feet are not red as in sladeni but vary 
from buffy to whitish. (2) The tail may be agouti or ochraceous to 
buffy but not white like that of haringtoni and its distal portion is 
not red like that of sladeni. (3) The dorsal pelage may be agouti or 
may vary about Ochraceous-Buff (XV) in those intergrades with 
haringtoni which have been called careyi, but differs in being thus 
tonally much darker than haringtoni, the dorsal pelage of which 
varies about Pale Ochraceous-Buff. 

Discussion. — The type specimen of hartoni is actually intermedi- 
ate between what is recognizable as the subspecies hartoni and sla- 
deni, possessing a red tail as it does. The geographic area occupied 
by hartoni, described in the discussion of sladeni as a "band" about 
the range of haringtoni, may be a very thin band along the Uyu 
River. Conceivably, the Uyu may for part of its mid-length actually 
separate haringtoni directly from sladeni. Should further collecting 
prove this to be the case, the subspecies hartoni as we recognize it 
here will then have to be redefined as two separate subspecies, one 
northeast of haringtoni, the other south of it. See the discussion of 
subspecies hartoni in the account of subspecies sladeni. 

Callosciurus erythraeus haringtoni (Thomas) 

Sciurus haringtoni Thomas, 1905, Ann. Mag. Nat. Hist. (ser. 7), 16, p. 314. 
Callosciurus solutus Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, p. 199. 

Types. — Sciurus haringtoni, BM No., juvenile male, from 
Maungkan, 25° N., 420 feet, upper Chindwin River, north Burma, 
collected December 14, 1904, by H. H. Harington; solutus, BM 
No., male (no skull), from Homalin, upper Chindwin River, 
north Burma, collected January 2, 1905, by H. H. Harington. 

Material examined, all from the east bank of the Chindwin River, 
Burma.— Hulaung (AMNH), four; Maungkan (AMNH), 20 topo- 


types, (BM), five; Kawya (AMNH), 12; Homalin (AMNH), 11; 
(BM), eight, (CNHM), two. 

Original description. — The type description applies so well in this 
subspecies to subsequently obtained material that we quote at length 
from it: "General colour of upper [pelage] 'cream -buff' along the 
dorsal area, the buff fading out on the sides, which are dull whitish. 
Individually the hairs of the back are whitish grey basally, with a 
broad cream-buff subterminal band and a minute black point. Head 
creamy whitish, with a slight buffy suffusion on the crown; the cheeks 
dull white. Ears whitish, both outside and in. [Ventral pelage], 
from chin to anus, bright sharply contrasted ochraceous buff (in the 
type; the second specimen nearer tawny ochraceous). Lateral line 
of demarcation [between dorsal and ventral pelage] very sharply 
marked in both specimens, and in the type emphasized by a black- 
ish line which runs from the middle of the front of the forearm, across 
the shoulders, down the sides and legs to the back of the heel. Fore 
limbs on outer side above this line of demarcation creamy whitish, 
like the flanks; beyond it, including the hands and the whole of the 
inner aspect, ochraceous buffy like the belly, or slightly paler. Back 
of upper part of hind leg whitish like body; inner side, ankles, and 
feet buffy like belly, rather paler on the digits. Tail above and below 
creamy buff proximally (the extreme tips of the hairs blackish), 
lightening to white terminally." 

Discussion. — In the large American Museum series only eight have 
the black lines which demark separation between the dorsal and ven- 
tral pelage really strongly developed ; 29 of the specimens have them 
weak and indistinct; and in 12 the lines are absent. All these speci- 
mens were collected between March 20 and 27, and this does not ap- 
pear to support Thomas's prediction that the presence or absence of 
the black line is likely to prove to be a seasonal phenomenon. The 
ventral pelage varies extremely in color from one individual to another 
in this subspecies (the color of the different parts of the ventral pel- 
age is remarkably uniform in any one individual), and the color of the 
dorsal pelage of the feet follows the variation of the ventral pelage 
in each individual. 

Habitat.— G. C. Shortridge {in Wroughton, 1916b, p. 293) com- 
ments as follows on the ecology of the vicinity of Homalin inhabited 
by C. e. haringtoni: "On the East bank. The jungle here changes 
completely and appears to be a curious kind of rather open, not very 
high, deciduous jungle with stretches of "kaing" grass. Country 
flat and swampy ..." 


Morris (1935, p. 667) gives more details from McCann's field 
notes. Between Hulaung and Maungkan, "All along the eastern 
bank were a number of beautiful Bauhinia trees in flower." And 
at Homalin, more extensive observations, "The forest on the east 
bank is composed of small deciduous trees among which are Careya 
arhorea, Holarrhena antidysenterica, Feronia, Dillenia pentagyna, Zizy- 
phus, Ficus glomerata and other species of figs, also Congea, Bomhax, 
Comhretum, Elaeocarpus and a large species of Smilax which was in 
flower. ... In the evergreen patches bordering the reservoir and 
stream the following were noticed. . . . The deciduous forest con- 
tained large numbers of a species of Erythrina ..." 

Callosciurus flavimanus (Geoffroy St. Hillaire) 

Definition. — The species Callosciurus flavimanus includes the sub- 
species quinquestriatus, gordoni, shanicus, michianus, zimmeensis, 
thai, atrodorsalis, siamensis, pranis, rubeculus, hendeei, castaneoven- 
tris, flavimanus, griseimanus, gloveri, bonhotei, styani, ningpoensis, 
and thaiwanensis and named forms here included in these subspecies. 
The species flavimanus occurs throughout the Indochinese Subregion 
east of the Sittang and upper Irrawaddy rivers (see fig. 13), excepting 
the areas of Thailand and Cambodia and part of Laos along the 
Mekong, where the species finlaysoni occurs. 

Diagnosis. — In this extremely variable species no pelage or other 
characters have been found which distinguish all of its subspecies 
from all of the subspecies of its closest relatives. Table 6 provides 
body and skull dimensions of some types included in this species. 

Relationships to other species. — This discussion evaluates the evi- 
dence for recognizing the species Callosciurus flavimanus to have the 
geographic range, and to include the subspecies shown in fig. 13. 
To follow it the reader will need to make constant reference to that 
figure and possibly some reference to the Classification provided near 
the beginning of the paper. Inasmuch as great rivers are seen to be 
barriers to tree squirrels in the Indochinese Subregion, the most 
likely place to look for evidence of intergradation between Callosciu- 
rus erythraeus west of the Irrawaddy River with some form of its 
closest relative across the river would be as far up the river as the 
squirrels occur. There the narrower river would be less of a barrier, 
and it seemed fair to ask whether even at 28° latitude the elevation 
might permit ice to bridge the river in enough places often enough 
to keep up genetic exchange between the populations on opposite 
sides of the river. Lord Cranbrook, who collected birds and mam- 


mals in the Nmai Hka headwaters of the Irrawaddy in 1931, com- 
ments (letter of May 5, 1961) on this: "The Nam Tamai and Adung 
are both headwaters of the Nmai Hka, and there were no snow 
bridges until about 7000 or 8000 feet on the Adung, at which point 
squirrels could have crossed on trees." He reached the Adung Val- 
ley on January 25th and collected in it about Tahawndam until May 
when the snow melted back enough to let his expedition go farther 
up the valley (Cranbrook, in Kinnear, 1934, p. 348). Some of Lord 
Cranbrook's published observations (loc. cit.) bear on the present 
problem, "On the fourth day the [Adung] valley turned to the north 
and there seemed to be a definite change from typical subtropical 
hill jungle to forest of a more temperate type. Conifers were quite 
numerous in the valley bottom. . . . Even the fauna seemed to 
change, as above this bend I saw none of the red-bellied Callosciurus 
erythraeus which were so plentiful in the . . . lower Adung, and the 
long-nosed squirrels, Dremomys pernyi and D. [lokria] seemed to take 
their place. The next day we arrived at Tahawndam ..." Since 
the February and March records of bird specimens (Cranbrook in 
Kinnear, 1934) are consistently recorded as 6000 feet, this is surely 
the elevation at Tahawndam. In Lord Crankbrook's experience, 
therefore, the red-bellied Callosciurus erythraeus did not range up to 
the elevation at which snow bridges occurred. 

On having seen the above arrangement and implications of his 
remarks. Lord Cranbrook (letter of June 25, 1961) expressed doubt 
that the Adung River is a barrier to squirrels even below 6000 feet. 
He then adds: "I think, though, that it is reasonable to assume that 
the climate in the Adung Valley has become less rigorous fairly re- 
cently. The glacier from which the river rises was obviously in 
retreat, with a long, convex snout, and the appearance of the val- 
ley below it made Kingdon Ward think that the retreat had been 
pretty rapid. The glaciers from which the Taron rises, the other 
side of the Namni La, were also in retreat and that valley too showed 
signs of recent ice action. If there had been much more ice and 
snow at the head of the comparatively short valley of the Adung 
and also at the head of the even shorter valley of the Seingku, the 
Callosciurus-Dremomys boundary . . . might well have been pushed 
further south to a point where the Tamai was in fact a barrier." 

Downstream C. erythraeus sladeni opposes the forms C. flavi- 
manus gordoni and C. /. shanicus across the Irrawaddy River, and 
there is no hint from the material reported herein from American 
museums, of direct genetic exchange between sladeni and either of 



these other two. To the north of sladeni the subspecies C. e. inter- 
medius occupies the region about 200 miles wide between the Brah- 
maputra's north-south course and the Irrawaddy's eastern fork the 
Nmai Hka. Opposite intermedius across the Nmai Hka is quinque- 
striatus, which has stood as a separate species because of its extra- 
ordinary ventral black and white stripes (Ellerman and Morrison- 
Scott, 1951, p. 488; Anthony, 1941, p. 89). Since quinquestriatus is 
shown below to intergrade with gordoni on the south, it becomes im- 
portant to consider whether quinquestriatus may also intergrade with 
intermedius near the headwaters of the Nmai Hka, or if there is in 
fact a species level difference between the last two. 

The 1938-39 expedition from the American Museum of Natural 
History to the mountain range between the Nmai Hka and the Sal- 
ween River, remained south of 26° 30 'N. They collected a substan- 
tial series of quinquestriatus (Anthony, 1941, p. 90), but there the 
elevation of the barrier river Nmai is apparently less than 700 feet. 
This is at least 100 miles south of the area where intergradation 
would be most likely to take place. The area between the barriers 
of the Nmai and Sal ween is less than 40 miles wide and suitable 
habitat for tropical tree squirrels is probably narrowed to somewhat 
less than that. If other influences for differentiation are the same, 
one could expect perhaps even greater differences to occur in this 
narrow 100-mile strip north of the northernmost available specimen 
of quinquestriatus than are now known, and shown below, to occur 
in the first 100 miles south of the range of quinquestriatus. (See 
fig. 13.) 

The similarities in color characteristics of the pelage, then, be- 
tween American Museum of Natural History material of intermedius 
and quinquestriatus are: 1. Generally agouti dorsal pelage of about 
Dresden Brown (with a reddish suffusion on the middorsum bringing 
it to about Cinnamon Brown there) over the head, back, tail, sides, 
legs, and feet. 2. The digits and sometimes the feet are blacker 

Fig. 13. The geographic distribution of two species, the red-bellied squirrel, 
Callosciurus erythraeus, A to G, and the belly-banded squirrel, Callosciurus flavi- 
manus, H to Z, as revealed by material examined (except for open dots on Formosa 
which indicate records from literature). Subspecies of erythraeus (west of the 
Irrawaddy): A, erythraeus; B, erythrogaster; C, bhutanensis; D, intermedius; E, sla- 
deni; F, bartoni (triangles); G, haringtoni (squares). Subspecies of flavimanus 
(east of the Irrawaddy-Sittang Valley): H, quinquestriatus; I, gordoni; J, shanicus; 
K, michianus; L, zimmeensis; M, thai; N, atrodorsalis; O, siamensis; P, pranis; 
Q, rubeculus; R, hendeei; S, castaneoventris: T, flavimanus; U, griseimxinus; V, glo- 
veri; W, bonhotei; X, styani; Y, ningpoensis; Z, thaiwanensis. 



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agouti than the general dorsal pelage. 3. The agouti guard hairs 
of the dorsal body pelage have from one to three light bands, gen- 
erally two, but on some individuals three is common. 4. The tail 
in ventral view appears scarcely more distichous than in dorsal. 
5. There is no pronounced eye ring. 

The distinctions in color of the pelage between intermedius and 
quinquestriatus are: 1. In five of the six intermedius the venter is en- 
tirely red (not bisected by a band of agouti), but in quinquestriatus 
the most extreme condition of bisection in the whole subregion oc- 
curs. (The bisecting band is black instead of agouti, the divided 
ventral pelage is white instead of red, and the outer margins of the 
white are bordered with black.) 2. C e. intermedium has no black 
tip to the tail, but quinquestriatus generally has a large and conspic- 
uous black tip to the tail (which becomes subterminal in some speci- 
mens wherein the black terminal hairs are generously tipped with 
brown). 3. The hairs of the tail of intermedium have five blackish 
bands (additional to a short, black tip), but those of quinquestriatus 
have only four. 4. The ears of intermedium are no redder than the 
pelage of the middorsum, but in quinquestriatus the ears are redder 
than the middorsum (except in the very red specimens from low ele- 
vations across the Irrawaddy from Myitkyina). 5. The rostrum and 
the face below and slightly encircling the eye of intermedium is a 
noticeably cooler gray than the nape, but in quinquestriatus these 
parts are huffier and warmer in color than the nape (again excepting 
the very red lowland specimens). 

The above differences between these two forms are much greater 
than those between quinquestriatum and gordoni and even somewhat 
greater than those between quinquestriatus and shanicus, the next 
form south of gordoni. Since the differences between quinquestriatus 
and gordoni and between gordoni and shanicus are in both cases very 
great ones for subspecies in the subgenus Callosciurum of this sub- 
region, it appears that the differences between quinquestriatum and 
intermedium are too great and the similarities too general for one to 
presume their relationship to be subspecific. 

Since on the basis of degree of difference, forms intermedium and 
quinquestriatus evidently represent two species, one should note addi- 
tionally that a geographic intensification of differences between the 
two species progresses toward the probable point of most frequent 
contact. In the eastern species, for which flavimanum is the oldest 
acceptable name, the pelage character which may be called bisected 
venter is one of the most widespread in the Oriental Region (south 


for 1700 miles through six subspecies, east for 1500 miles through 
five subspecies). In gordoni (not far from the range of intermedius) 
it intensifies to its strongest agouti expression. In quinquestriatus, 
closer to the range of intermedius, it becomes stronger still by turn- 
ing to black. Approaching from the south through subspecies ruhec- 
ulus, pranis, siamensis, atrodorsalis, thai, zimmeensis and gordoni, or 
from the east through thaiwanensis, castaneoventris, hendeei, and 
michianus one finds the rich red venter (divided or undivided) . This 
is similar to the venter of C. erythraeus intermedius (and all but one 
form of erythraeus), but upon closest approach to the probable point 
of latest, or most frequent species contact, in C. flavimanus quinque- 
striatus the red venter is lost and replaced with black and white 
stripes. This provides extreme contast with the bright red venter 
of intermedius. Approaching from the west, in C e. hhutanensis one 
finds the black tail tip, reddish face, reddish ear tips, and divided 
venter that occur in eastern subspecies of the species flavimanus. 
Moving closer to the probable point of most frequent, or recent, con- 
tact between the species, one finds all of these characteristics gone 
or greatly reduced in intermedius. 

The above phenomenon appears to be very similar to what has 
been called "character displacement" (Brown and Wilson, 1956), 
differing from their definition of character displacement in that there 
is now no known zone of overlap of ranges of the two species. 

It is apparent on examination of the range map (fig. 16) of Cal- 
losciurus pygerythrus that the former river course of the Irrawaddy- 
Sittang must have been the eastern barrier to spread by that species, 
and for the most part, still is today. The Nmai Hka branch of the 
upper Irrawaddy apparently still is the barrier to eastward spread of 
pygerythrus in the region where the species pygerythrus is sympatric 
with C. e. intermedius. Although the Nmai is not shown so large on 
maps as the Salween, Mekong, or Brahmaputra, it seems likely on the 
basis of squirrel distribution that it was larger in the past and sub- 
sequently lost some of its headwaters by stream piracy. Such a 
former condition would better explain why the Irrawaddy shows so 
much evidence of having provided the greatest barrier to east and 
west spread of tree squirrels in the Indochinese Subregion. 

Callosciurus flavimanus quinquestriatus (Anderson) 

Sciurus quinquestriatus Anderson, 1871, Proc. Zool. Soc. London, 1871, p. 142. 
Sciurus beebei J. A. Allen, 1911, Bull. Amer. Mus. Nat. Hist., 30, p. 338. 


Callosciurus quinquestriatus imarius Thomas, 1926, Ann. Mag. Nat. Hist. 

(ser. 9), 17, p. 640. 
Callosciurus quinquestriatus Sylvester Thomas, 1926, Ann. Mag. Nat. Hist. 

(ser. 9), 17, p. 641. 

Types. — Sciurus quinquestriatus, IM No. 10163, specimen from 
Ponsee, east of Bhamo, Burma, in the Kakhyen Hills, taken March 4, 
1868, by John Anderson; beebei, AMNH No. 32624, adult male, 
originally described in error as from Sarawak, but fixed by Goodwin 
(1953, p. 282) as Sansi Gorge, Burma-Yunnan boundary, Decem- 
ber 7, 1910, collected by C. William Beebe; imarius, BM No. 20.8.- 
7.6, old male, from west flank of Imaw Bum, Kachin, Burma, lati- 
tude 26°10'N., longitude 98°30'E., October 21, 1919, by F. K. 
Ward; Sylvester, BM No., adult male (skull lost), from the 
forests of the Shweli-Salween divide, 9000 feet, west Yunnan, China, 
collected September, 1925, by G. Forrest. 

Material examined, from northeastern Burma. — 15 miles north of 
Myitkyina on east side of Irrawaddy River (USNM), eight; six miles 
north of Myitkyina (USNM), one; Porg-ka-tong (AMNH), one; 
Pum-ka-tong (AMNH), one; between Nomoyoung and Seniku 
(AMNH), one; between Tamu and Chipwi (AMNH), three; be- 
tween Chipwi and Laukhauna (AMNH), two; between Tamu and 
Tanga (AMNH), two; Laukhaung (AMNH), four; Pyepat (AMNH), 
three; between Pyepat and Langyang (AMNH), two; above Tsonma, 
8300 feet (AMNH), one; Gangfang, 5200 feet (AMNH), one; Hkam- 
kawn, 4000 feet (AMNH), one; Hpimaw, 7600 feet (AMNH), two; 
Htawgaw (AMNH), one; Sima, Myitkyina, 4500 feet (BM), three; 
Sui Kin, Bhamo (BM), three; Teinzo, northeast of Bhamo (BM), 

Pelage color. — Description of Indian Museum Paratype, a stuffed 
study skin examined in January of 1960: Dorsal pelage agouti of 
about Cinnamon Brown (XV) on crown, nape and back, but paling 
on tail and sides to Dresden Brown. Ears in part redder, about 
Ochraceous Tawny. Feet like back but digits blacker. Tail hairs 
with blackish tips plus 4 blackish bands, but tip of tail light, about 
Buckthorn Brown. Agouti dorsal pelage has two distinct light bands 
and black on the tips and between light bands. Ventral pelage has 
two longitudinal light stripes. Light Ochraceous Buff from posterior 
end of throat to anus, separated and bordered by blackish bands 
(thus making two "white" and 3 "black" stripes) of the same length. 
The light bands are about 10 mm. wide on the thorax but 5 mm. or 
less for most of their length. The blackish middle stripe is similarly 


wide but the two lateral ones seem narrower. The middle stripe is 
definitely agouti on the thorax. The "black" stripes seem faded and 
are only about Prouts Brown. The hind legs are agouti all the 
way around, definitely, but blackish pelage apparently extends out 
the ventral surfaces of the fore limbs. The tail has a subterminal 
black tip. 

In the original description Anderson recorded the light stripes as 
"pure white" and their laterally bordering stripes as black and this 
black as "expanding on the inside of the thighs." Probably during 
the intervening 90 years the white has become less white and the 
black less black, for the American Museum Natural History material 
agrees emphatically with Anderson's description in these three par- 

Discussion. — At the very beginning of knowledge of this squirrel, 
Anderson (1871, p. 142) considered it most closely related to [Callo- 
sciurus flavimanus] gordoni and included the two kinds under one 
common name, "the belly-banded squirrels." Nevertheless, the 
uniqueness of the five vividly contrasting black and white longitu- 
dinal bands of quinquestriatus has evidently discouraged all subse- 
quent students down to the present from considering that these two 
squirrels may be conspecific. There is, however, evidence that their 
ventral markings possess a common pattern which suggests that the 
amount of genetic difference between quinquestriatus and gordoni is 
not very great. 

The following observations are the basis upon which quinquestri- 
atus and gordoni are here considered conspecific (see fig. 13). The 
gordoni specimen CNHM No. 32519 from 27 miles north of Bhamo 
has a blackish midventral stripe and incipient black showing in the 
edge of the agouti of the animal's side where the agouti meets the 
red of the venter. In gordoni specimen AMNH No. 43217 from Hui 
Yao, Yunnan, the edges of the 7 mm. wide midventral agouti stripe, 
as well as the ventral edges of the agouti sides of the animal, are 
blackened. AMNH No. 43217 represents no great departure from 
the characteristics of gordoni, but its midventral stripe is precisely 
like that of many of the American Museum specimens of quinquestri- 
atus. Even the ventral edges of the sides of AMNH No. 43217 
would need to become but little blacker to equal the blackness of 
some examples of American Museum material of quinquestriatus. 
Especially significant among the large series of quinquestriatus is 
AMNH No. 114907 from between Tomu and Chipwi, Burma. This 
specimen although preponderantly quinquestriatus in character ap- 


preaches the characteristics of gordoni in several ways: (1) The mid- 
ventral stripe is quite like that described for AMNH No. 43217, and 
the ventral edges of the sides are only a little blacker. (2) The sub- 
terminal black tip to the tail is short and rather subdued. (3) The 
two ventral stripes which are ordinarily white in quinquestriatiLs, are 
Apricot Buff (XIV). 

The ventral pelage of gordoni, thus, may be said to be three- 
striped, or to consist of a 6 or 7 mm. agouti longitudinal mid-stripe 
which separates the red of the remainder of the venter into two 
broader, rather stripe-like, sections. In quinquestriatus the midven- 
tral stripe is blackened on its edges or sometimes is entirely black. 
The outer edges of the remaining ventral pelage is black, making two 
lateral black stripes of a width approximating that of the middle 
stripe, but including also the entire ventral pelage of the limbs. The 
remaining two areas of ventral pelage, generally about ten to twelve 
millimeters wide, are white. Thus, there are characteristically two 
white stripes intersticed between three black ones. 

The dorsal pelage of these two forms differs in but small details. 
Dorsal pelage of both body and tail is an agouti varying about Buffy 
Brown (XL) in quinquestriatus but is generally less red in gordoni. 
The subterminal black tip of the tail is generally longer in quinque- 

No evidence of intergradation between C /. quinquestriatus and 
subspecies [C. erythraeus] intermedius or [C. flavimanus] michianus 
has reached our attention as such. Probably the Sal ween River is 
barrier enough to prevent quinquestriatus from intergrading with 
michianus (see fig. 13). It would seem surprising, however, if the 
Nmai Hka or its northeastern tributary, the Taron River, would long 
prevent C. e. intermedius from coming into contact with C. f. quin- 
questriatus. See discussion above in the account of the species flavi- 

Callosciurus flavimanus gordoni (Anderson) 

Sciurus gordoni Anderson, 1871, Proc. Zool. Soc. London, 1871, p. 140. 

Type, here designated. — IM No. 9268 a parous female taken at 
Bhamo, upper Burma, on February 24, 1868, by John Anderson. 
This specimen is here selected from the two cotypes as the type of 
gordoni because it is adult (whereas the other is not) and is rather 
typical of the subspecies in ventral color (whereas the other is not) . 

Material examined, all from a small area of northeast Burma 
and adjacent Yunnan. — Homoshu Pass, 8000 feet (AMNH), one, 


(CNHM), one, (MCZ), one; Taipingpu, 7000 feet, Shweli River 
(AMNH), two, (CNHM), one, (MCZ), one; "Hui Yao," 5500 feet, 
Tengyueh (AMNH), one; Watien, Tengyueh (AMNH), two; "Yun- 
nan" (AMNH), one; Kyu Loi, S. Shan States (ANSP), one; Shweli- 
Salween Divide, 7000-10,000 feet (CNHM), two, (BM), twelve; 
Bhamo, Burma (USNM), one topotype; Momauk, nine miles east 
of Bhamo (USNM), one; Cuchi, Yunnan (USNM), one; "Linchi- 
apu," Yunnan (USNM), one; Shweli Valley, 9000 feet (BM), two, 
8000 feet (BM), eight, 7000 feet (BM), one; hills northwest of 
Tengyueh, Yunnan (BM), three; "Watan," 5000 feet, Bhamo (BM), 
five; Machangkai, 6500 feet, Tengyueh (BM), one. 

Type description. — A description of the type from examination of 
it in 1959, follows: Ventral pelage Cinnamon Rufous (XIV), but 
divided all the way down the middle by an agouti stripe 3 mm. to 
5 mm. wide. The color of the venter extends over half the length 
of the throat. The remainder and muzzle are about Isabella Color, 
and so are circular areas around the four teats (about 10 mm. diam- 
eter). Dorsal pelage agouti, paler than Citrine Drab (XL). The 
sides nicely Dark Olive Buff (XL). Dorsal guard hairs have two 
buffy bands each 1^/^ mm. long separated by a black band 23^ mm. 
long. Tips are black. Head, legs, and feet like back. Ears like 
muzzle. Tail color like dorsum both dorsally and ventrally. Not 
perceptibly distichous below. Tail hairs have black tips and four 
black bands 3 mm. to 4 mm. long (and four 2 mm. to 3 mm. light 
bands and a shorter light base) . Tail has short blackish pencil end- 
ing in buff tip. Notably, we think, in both cotypes the mid ventral 
agouti stripe is most intensely agouti in the stripe's midlength rather 
than near the throat and in the present specimen, does not reach 
the throat. 

Discussion. — This rather well-marked subspecies appears from 
the material examined to occupy a very small range in northeastern 
Burma between the Irrawaddy River at Bhamo and the Salween 
River. Anderson (1879, p. 240) gives an impression of a somewhat 
greater range when he says, "I have collected about twenty-five 
specimens of this squirrel from various parts of Burma, north of the 
capital [Mandalay], . . ." However, it seems doubtful that gordoni 
occurs south of the Shweli River in Burma excepting as an intergrade 
with C. f. shanicus, for shanicus is known from several localities north- 
east of Mandalay along the road and railroad to Lashio. 

To the north gordoni appears to be replaced rather abruptly by 
quinquestriatus, and evidence of intergradation between the two has 


been described above under the discussion of subspecies quinquestri- 
atus. Subspecies gordoni differs from michianus and zimmeensis to 
the east in lacking the incipient black mark on the posterior two- 
thirds of the middorsum and in possessing a median agouti band 
which completely bisects the red pelage and which is itself the color 
of the agouti dorsum. To the south it seems most logical that gor- 
doni intergrades with shanicus from which it differs by possessing: 
(1) divided red ventral pelage, (2) a face which is just as dark agouti 
as its dorsum, and (3) a middorsum which lacks the incipient black 
on its posterior two-thirds. From subspecies [C. erythraeus] sladeni 
to the west across the Irrawaddy gordoni differs (so greatly that it is 
no wonder they have been considered separate species) by possess- 
ing: (1) the red venter divided, (2) the feet and rostrum agouti in- 
stead of red like the venter, (3) the tail colored like the back instead 
of extensively red-tipped. Details of the possibility of intergrada- 
tion with phayrei are considered under the species phayrei. 

Callosciurus flavimanus shanicus (Ryley) 

Sciurus atrodorsalis shanicus Ryley, 1914, Jour. Bombay Nat. Hist. Soc, 22, 

p. 662. 
(?) Sciurus griseopectus Blyth, 1847, J, Asiat. Soc. Bengal, 16, p. 873. 

Types. — Sciurus a. shanicus, BM No., an adult male 
from Gokteik, 2133 feet. North Shan States, Burma, taken April 23, 
1913, by G. C. Shortridge; griseopectus, IM No. 9369, a parous fe- 
male captive from unknown locality, presented in 1847 by Rajah R. 

Material examined, all from Burma. — Maymyo, 800 meters 
(AMNH), four, (BM), six; Gokteik, 2133 feet (BM), two, (CNHM), 
two; Pyaunggaung, 2794 feet (BM), four, (CNHM), one; Kawlaw, 
4400 feet, west of Taunggi (BM), one; northeast of Hoiparo, N, Shan 
States (BM), one; Mongha, N. Shan States (BM), one. 

Discussion. — Specimens of shanicus were taken at precisely the 
same elevations as pharyrei at three of the same localities (Maymyo, 
Gokteik, and Pyaunggaung). At Maymyo in 1937 Gerd Heinrich 
collected six phayrei between November 27 and December 25 and a 
series of four shanicus between December 3 and 20, He took one 
specimen of each form on one of these days (the 20th), Heinrich's 
four phayrei with substantial toothwear have head-and-body length 
ranging from 222 to 238 mm. Although three of the shanicus have 
substantial toothwear, all have head-and-body length between 200 
and 211 mm. These data are quantitatively weak but no indication 


is seen in them that shanicus is a seasonal or other color phase of 
phayrei. This and the many differences reported in the account of 
species phayrei suggest strongly that they belong to separate species. 
The form shanicus closely approximates gordoni in size and will 
doubtless be shown to intergrade with gordoni. The color charac- 
teristics of shanicus show evidence of relationship also to atrodorsalis 
and zimmeensis, but the Salween River probably prevents intergra- 
dation. See also comparisons made under C. p. janetta. 

Pelage color. — The two topotypes at Chicago have dorsal pelage 
of body, legs, feet, head, and tail Grayish Olive and incipient black 
shows on the posterior two-thirds of the middorsum on both. The 
abrupt, brief, black tip of tail has whitish tips. The ventral pelage 
is Old Gold (XVI) in color, and all of it is agouti. The Chicago Nat- 
ural History Museum specimen from Pyaunggaung agrees with this, 
and so do the four specimens in the American Museum of Natural 
History from Maymyo. 

//a6*7s.— Shortridge's field notes on this subspecies follow: "The 
most plentiful squirrel in Hsipaw State and around Maymyo, recall- 
ing Funamhulus palmarum of India in its habit of collecting around 
gungalows. The black mark on the back was seldom conspicuous 
except in immature specimens." (G. C. Shortridge in K. V. Ryley, 
1914, p. 721.) 

Type description. — Examination of the type of Sciurus griseopec- 
tus Blyth, 1847, reveals (in 1959) the following characteristics: Ven- 
tral pelage Ochraceous-Tawny to Tawny except for agouti throat 
and a midventral extension from the throat posteriorly halfway to 
the anus. Throat is Deep Olive Buff (XL). Muzzle and chin are 
Chamois (XXX), and so is a circular area 10 mm. diameter around 
each teat. Proximally the tail is lighter beneath. Dark Olive Buff 
(XL), than above. Deep Olive. Dorsal pelage agouti and entirely 
about Buffy Brown (XL). The ears are like the muzzle. Chamois. 
The dorsal guard hairs have two buffy bands each two mm. long and 
separated by a 2 or 3 mm. blackish band, and have black tips. The 
tail hairs are brownish buffy tipped and have four black bands. 
The tail's dull color may be in part due to the hair being old, but 
probably when fresh it is only a little less dull. The feet look faintly 
paler than the middorsum, but not paler than the sides. There are 
barely evident eye rings tending toward Chamois. The tail is not 
perceptibly distichous below. 

Discussion. — In the original description of griseopectus, Blyth 
(1847) mentioned two points important to identification of the re- 


gion whence the type must have come: the divided red venter, and 
"the tail being slightly black-tipped, but with pale ends to the hairs." 
These conditions occur in ningpoensis, michianus, and zimmeensis, 
as will be shown below, and in gordoni as reported above. Strong 
tendency to feet blacker than dorsum mitigates against all of these 
but ningpoensis. Anderson (1879, p. 239) made something of a case 
for the type of griseopectus having come from China, and placed it 
in the synonymy of castaneoventris Gray 1842. As restricted here, 
castaneoventris has feet too black and tail tip too different. The 
proper color relationship of foot to dorsum is common in the large 
AMNH series of ningpoensis, but in ningpoensis the muzzle and chin 
are rather consistently cool gray, and the tail hairs have five or even 
six blackish bands instead of four. In ningpoensis also three light 
bands on a guard hair in the pelage of the dorsum is common, where- 
as in the type of griseopectus the highest number noted was two. In 
conclusion, the probability of the type specimen of griseopectus be- 
ing from China east of the Sal ween River seems slight to us, even 
though the sample of ningpoensis is so variable, and even though the 
area of southern China from which no material representing this spe- 
cies has been available, is extraordinarily large. 

When material becomes available from an area of intergradation 
between gordoni and shanicus, it seems likely that individuals quite 
like the type of griseopectus may be found among them. The ven- 
tral pelage color would be quite intermediate, and shanicus could 
contribute the proper muzzle, chin, ear, and foot color and the correct 
number of blackish bands on tail hairs and light bands on guard hairs 
of the dorsal pelage. The favorable gordoni influence would be toward 
browner dorsal pelage and less distichous tail shape. At present 
writing this identification of griseopectus seems more probable than 
geographic location in eastern China. Until someone can identify 
material from known localities with the type of griseopectus, however, 
it seems best to let the name shanicus stand and to associate grise- 
opectus with it only uncertainly. 

Callosciurus flavimanus hyperythrus (Blyth) 

Sciurus hyperythrus Blyth, 1855, Jour. Asiatic Soc. Bengal, 24, p. 474. 

Type. — IM No. 9478, an individual from Tenasserim, presented 
in 1852 by Major Berdmore. 

Ellerman and Morrison-Scott (1951, p. 479) suggest the Sittang 
Valley as the geographic source of the type. 


Type description. — The following characteristics of the type were 
observed by one of us in 1959: Ventral pelage, including that on 
limbs, is entirely Bay or Claret Brown except for the chin which is 
infiltrated by agouti and is about Hazel. The dorsal pelage is agouti 
and is about Light Brownish Olive (XXX). This color is ubiquitous 
except on the ears and the head forward of the ears, where it merges 
into Hazel. The short pelage on the rather battered ears is clearly 
deeper red, about Kaiser Brown or even Hays Russet. There is no 
eye ring. Dorsal guard hairs generally have two light bands each 
somewhat exceeding one mm. in length and separated 3 or 4 mm. 
from each other. The tail hairs have five blackish and five light 
bands and their tips are blackish on the proximal half of the tail but 
reddish on the distal half. The tail shows a faint tendency to be 
distichous beneath. 

Discussion. — Material representing the species flavimanus from 
lower Burma is so scarce that although we are doubtful regarding 
whether the type specimen came from the vicinity of the town of 
Tenasserim or just from somewhere in the province of Tenasserim 
(which seems from usage to have extended north at least as far as 
the lower Sittang Valley), it is not possible for us to locate the most 
probable geographic origin of the type by comparing it with material 
from various parts of Tenasserim. In any case the type locality is 
certainly not Moulmein as questioningly suggested by Robinson and 
Kloss (1918, p. 199) unless they would wish to imply across the Sal- 
ween River, for Moulmein is designated type locality for C. f. atro- 
dorsalis and has proven to be quite acceptable as such. Martaban 
across the Salween would be a perfectly acceptable type locality, but 
not necessarily any better than the Sittang Valley 70 miles farther 
north. However, until specimens become available for study from 
these places it must be left a little uncertain where the type locality 
of this named form is, and if it is in fact a subspecies, where its whole 
range may be. Burma between the Sittang and the Salween seems 
the most reasonable possibility, but it remains to be proven. 

Callosciurus flavimanus michianus (Robinson and Wroughton) 

Sciurus castaneiventris michianus Robinson and Wroughton, 1911, Jour. Fed. 
Malay States Mus., 4, p. 234. 

Sciurus castaneoventris haemobaphes G. M. Allen, 1912, Proc. Biol. Soc. Wash- 
ington, 25, p. 177. 

Types. — Sciurus castaneiventris michianus, BM No., 
old female, from Mee Chee, Yunnan, collected January 5, 1903, by 


F. W. Styan; haemobaphes, MCZ No. 13693, adult male, from Chih- 
ping, southeast Yunnan, collected February 26, 1911 by Kobayashi. 

Material examined, from Yunnan, China. — "Ssu Shan Chang," 
9000 feet, Likiang (AMNH), three; Yunnanfu, 6400 feet (AMNH), 
two; 15 miles south of Kunming [= Yunnanfu] (USNM), one, 
(MNHN), one; Yunnanyi, 6500 feet (AMNH), one, (CNHM), one; 
Hangai, 5900 feet, 40 miles east of Talifu (AMNH), one; Chao Chu, 
6700 feet, south end of Lake Talifu (AMNH), one; Shan Kuang, 
6000 feet. Lake Talifu (AMNH), two, (CNHM), three, (MCZ), one; 
Mekong-Salween divide at 28° 20' N. latitude, 7000 feet (BM), one; 
Chepei, Fumin (AMNH), one; Kao Chiao, Yunnanfu (AMNH), 
six, (MCZ), one; Lungkai, Makaihsien (AMNH), six; Lungkai, 
Wutinghsien (AMNH), one; "Kun Yang" (AMNH), one; "Nda Mu 
Cho" (USNM), one; Makaihsien (MCZ), one; Likiang, 9000 feet 
(MCZ), three, (USNM), six, (CNHM), four, (BM), one; Chihping 
[Shihping] (MCZ), four; "Hofuping Mountains," Mekong Valley 
(USNM), three; Chiihsiung, 160 miles east of Yunnanfu (CNHM), 
one; Nui Kai, 7200 feet, 30 miles north of Talifu (CNHM), one; Lo 
Tu Kow [Lukou] (BM), one; six miles north of Luchang (BM), two. 

This subspecies seems to be bounded on the northeast by the 
Yangtze, but allocation of the Yung-pei-ting specimen to gloveri is 
made on this assumption in absence of adequate color notes on the 
specimen. The southwestern boundary of the range of michianus is 
probably the Mekong River. To the south in southern Yunnan 
michianus must surely intergrade broadly with Callosciurus flavi- 
manus hendeei. 

Diagnosis. — Subspecies michianus generally differs from gloveri to 
the north by possessing: (1) an agouti throat and a wedge from it 
extending posteriorly as an incipient midventral agouti stripe; (2) an 
incipient blackening of the pelage on the posterior two-thirds of the 
mid-dorsum; (3) a short, incipient, sub terminal, black tip to the tail; 
and (4) digits which are often notably blacker agouti than that of 
the animal's own gray dorsal pelage. Subspecies michianus differs 
from castaneoventris in characters (2) and (4) above, and in lacking 
a reddish infusion in the agouti dorsal pelage. From hendeei, michi- 
anus differs by characters (1) and (2) above, and the feet of hendeei 
are substantially blacker, and the black tail tip of hendeei is substan- 
tially blacker and longer, than that of michianus. Across the Me- 
kong River from C. flavimanus michianus is a subspecies, C. flavi- 
mauns gordoni, from which michianus differs only by being paler 
and larger. 


Callosciurus flavitnanus zimmeensis (Robinson and Wroughton) 

Callosciurus atrodorsalis zimmeensis Robinson and Wroughton, 1916, Jour. 

Fed. Malay States Mus., 7, p. 91. 
Callosciurus ferrugineus primus Allen and Coolidge, 1940, Bull. Mus. Comp. 

Zool., 87, p. 157. 

Types. — zimmeensis, BM No., an adult female taken 
at Chiengmai, 2100 feet, northern Siam on April 12, 1908, by T. H. 
Lyle; primus, MCZ No. 35352, old female from Mae Wan River, 
1500 feet, near Mount Souket, north Thailand. 

Material examined, from Yunnan, China. — Mucheng, 7000 feet 
(AMNH), four, (CNHM), two, (MCZ), one; Hsiaomengtung, 6000 
feet (AMNH), one; Tashutangh, 6000 feet (AMNH), one; Chenkang, 
3000 feet (AMNH), two; Chaunglung, Salween ferry, 2000 feet 
(AMNH), four, (CNHM), three, (MCZ), one; and Nam Ting River 
at Burma border (AMNH), four, (CNHM), two, (MCZ), two; 
Mengting, 1700 feet (AMNH), one. 

Material examined, from Burma. — Malipa (AMNH), one; Ban 
Sob Pa, Salween River (USNM), two. 

Material examined, from Thailand. — Doi Soutep (AMNH), three, 
(BM), seven, (USNM), one; Mt. Chieng Dao (AMNH), one, 
(USNM), three, (ANSP), three; Pa Kwe, Chiengmai (AMNH), 
two; Mt. Doi Pui, Chiengmai (AMNH), two; Mong Hawt, Chieng- 
mai (AMNH), two; Ban Kang, base of Doi Angka (USNM), two; 
Chomtong (USNM), one; Mekhan (USNM), one; Waterfall Me- 
kang (USNM), one; Mae Hong Song (BM), two, (USNM), one; 
Doi Hua Mot (USNM), one; Doi Angka (MCZ), 13, (CNHM), one; 
Chieng Mai (ANSP), two, (BM), four; Mee Tan [Ban Mae Tan 
Nua] (BM), one; Ban Meh Na (NR), two; Doi Par Sakeng (NR), 
three; Muang Pai, 600 meters (BM), one; "Muang Sang," 425 me- 
ters, north of Chiengmai (BM), one; Me Ping River at IS** 00' N., 
235 meters CBM), one; "Me Ping River" (BM), four. 

This subspecies occupies the waveringly wedge-shaped area be- 
tween the Salween River and the Mekong River from about 18° 
north latitude to about 25°, and possibly to 26° or beyond. The 
type locality is rather close to the area of intergradation with atro- 
dorsalis on the south. 

Diagnosis. — Subspecies zimmeensis differs from michianus on the 
east by being smaller and darker, from gordoni on the west by pos- 
session of the incipient black posterior middorsum and less nicety of 
the midsagittal division of the red venter, from shanicus by possess- 
ing a red, divided venter and five dark bands on the tail hairs, from 


hendeei by having at least incipient bisection of the ventral red pelage 
and by having the tail colored like the dorsum or with only a thin 
buffy tipping (in some southern material), and from atrodorsalis and 
thai by having the posterior two-thirds of the middorsum only in- 
cipiently black (i.e., without an area of pure black). 

Discussion. — Note in Figure 13 that zimmeensis material is widely 
divided into two lots. The difference between the northern (Tem- 
perate Zone) material of zimmeensis and its neighbor across the Me- 
kong is rather slight when compared with characters differentiating 
other subspecies of this revision of Callosciurus flavimanus. If this 
Temperate Zone sample of zimmeensis had shown greater differences 
from material from Thailand (300 miles into the tropics) we should 
have included the northern lot in michianus rather than naming it 
as a new subspecies. 

For discussion of the named form primiis see the account of 
C finlaysoni menamicits. 

It is interesting that about the Tropic of Cancer where the pop- 
ulations of species flavimanus have been sampled on both sides of 
the Salween and Mekong rivers which have run closely parallel to 
one another for about 400 miles, the samples from either side of the 
Salween are much more highly differentiated from each other than 
those from either side of the Mekong. See Table 7. 

Table 7. Subspecies of C. flavimanus separated by Salween River and Mekong 
Rivers 300 kilometers north and 300 south of the Tropic of Cancer. 

E. of Salween 

W. of Salween, E. of Mekong 

W. of Mekong 










Callosciurus flavimanus thai (Kloss) 

Sciurus atrodorsalis thai Kloss, 1917, Jour. Nat. Hist. Soc. Siam, 2, p. 285. 

Type. — Field No. 2474/CBK (not seen), from Raheng, central 
Siam, collected July 1916 by C. Boden Kloss. 

Material examined, from Thailand. — Mewong River, 1000 feet, 
40 miles east of Um Pang (AMNH), two; Mewong River, 800 feet, 53 
miles east of Um Pang (AMNH), two, (BM), four; 20 miles west of 
Kempenpet (AMNH), one, (BM), one; "Siam" (AMNH), one; 


Muang Kan Bouri [Kanchanaburi] (USNM), three; Ban Pong 
(USNM), one; Hue Yah Pla, 2500 feet (USNM), two; Me Taqua, 
800 feet (USNM), five, (CNHM), one; Wang Kien (USNM), one; 
Sai Yoke (BM), one; "Dooweeklo," 1500 feet, Mae Klong (BM), 
one; Mae Taw Forest, 900 feet, Raheng (BM), five; Lai Yoke [Ban 
Sai Yoke] (BM), one; 10 miles west of Raheng, 600 feet (BM), one; 
Raheng, 115 meters (BM), one; 28 miles east of Um Pang, 2000 feet 
(BM), two; Doi Mei Lai [Doi Mae Lai] (USNM), one; Doi Mae 
Kong Kha (USNM), two; Banphotphisai, Nakon Sawan Prov. 
(USNM), four; Slokbai [Ban Salok Bat], Kano, Kamphaeing Phet 
(USNM), one; "Tumphol," Kha Nu, Kamphaeing Phet (USNM), 
six; Ban Hua Thanon, Klong Klung, Kamphaeing Phet (USNM), six. 

Material examined, from Burma. — Lampha, Tenasserim (AMNH), 
one, (BM), three. 

The zimmeensis from northern Thailand show some intergrada- 
tion with the subspecies thai to the west and southwest, in occasional 
occurrence of some pure black pelage on the posterior two-thirds of 
the middorsum, and in the general occurrence of long light tips on the 
tail hairs which give to the posterior half of the tail an overlay of 
straw color (Pale Ochraceous Buff to Light Ochraceous Buff, XV). 

Diagnosis. — The subspecies thai is distinguished from zimmeensis, 
however, by having the following characteristics; (1) regular occur- 
rence of the solid black band of pelage 20-30 mm. wide along the 
posterior two- thirds of the mid-dorsum; (2) the rostrum, and occa- 
sionally the crown, is colored rusty (ranging from only Buckthorn 
Brown to Hazel) ; whereas in zimmeensis the rostrum and crown are 
always a cool gray. 

Callosciurus jiavimanus thai is distinguished from atrodorsalis by: 
(3) having its feet a blacker agouti than the legs; (4) having its hind- 
quarters not notably redder than its forequarters; (5) having long 
straw-colored tips to the tail hairs (Light Buff to Light Ochraceous 
Buff) instead of reddish (Ochraceous Buff to Cinnamon Rufous). 

Habits. — Callosciurus jiavimanus thai appears to occupy a rain 
shadow zone in Thailand and is probably found where the forest is 
primarily deciduous; whereas the following subspecies, atrodorsalis, 
apparently occurs where the forest is primarily broad-leafed ever- 
green rain forest. 

Callosciurus flavitnanus atrodorsalis (Gray) 

Sdurus atrodorsalis Gray, 1842, Ann. Mag. Nat. Hist. (ser. 1), 10, p. 263. 


Cotypes.—BM Nos. 41.1818 and 41.1819, both adult males re- 
ceived from the East India Company, "from Bhotan," corrected by 
Ryley (1914, p. 663) to Moulmein. 

Material examined, all from Burma. — Taok Plateau, 3050 feet 
(AMNH), eight, (BM) four; Zami R., 100 mi. S. of Moulmein (BM), 
one; Kokareet [Kawkareik], Tenasserim (USNM), one, (CNHM), 
one, (BM), three; Mt. Nwalabo, Tavoy District (BM), two; Doonsa 
(BM) one; Thaungyin Valley (BM), one; "Lathorgu," Myawadi 
(BM), one; Ban Sob Pa, Salween R. (USNM), two; Myawadi (BM), 
two; Sookli [Sukli], Myawaddy Rd. (BM), one; Maitho, Thaungyin 
River, Tenasserim (BM), one; Haungtharaw River (BM), one; 
"Shan Mepa," Amherst (BM), three. 

Pelage color. — One striking character that shows the relationship 
between the subspecies atrodorsalis and thai is, of course, the broad 
(20-30 mm.) band of pure black pelage on the posterior two-thirds 
of the mid-dorsum. Many subspecies of flavimanus have an incip- 
ient manifestation of the black band but only atrodorsalis and thai 
have it pure black. 

The agouti dorsal pelage varies considerably in color individually 
among specimens from the rainforest of the Tenasserim region, de- 
pending upon the amount of infusion with red. The rostrum in the 
atrodorsalis at the American Museum of Natural History is redder 
(Hazel to Kaiser Brown, XIV) than the dorsal pelage on the same 
individual. The anterior dorsal pelage in the Tenasserim AMNH 
series ranges from about Hazel in the reddest to about light Brown- 
ish Olive (XXX) in the grayest. The posterior half of the dorsal 
pelage (exclusive of the black stripe) is redder on each individual 
than the anterior half, the range in the same series being about 
Russet (XV) to Saccardo's Umber (XXIX). The tail is just about 
as reddish as the hindquarters on each individual. The feet, unusual 
in this species, are no blacker agouti than the legs. 

Discussion. — This subspecies occupies the very heavy rainfall area 
of Tenasserim and is much redder than the adjacent subspecies thai 
which occupies the contiguous rainshadow area in Thailand. 

Anderson (1879, p. 233) placed a great deal of stress on the oc- 
currence of two color phases in atrodorsalis, the more common one 
with a broad black band on the back, the other without. He found 
that the difference between the two was not sexual dimorphism and 
that neither color phase exclusively represents the stage of maturity. 
Whether it is seasonal, his material was not adequate to reveal, and 


we have examined too few of the ones lacking black patches to say. 
For the geographic range that Anderson attributed to atrodorsalis 
(Moulmein to Malacca), the difference has now been shown to be 
geographic, indeed, but he was also concerned with the local varia- 
tion in the Moulmein area and to the east of Moulmein. Here he 
declared the evidence was non-geographic but he did not state the 
evidence in adequate detail for one confidently to agree. Anderson 
(1879, p. 235) also states that "atrodorsalis is very common in Marta- 
ban. ..." We have seen no specimens from there, and since neither 
caniceps nor phayrei are known to have crossed the Salween River, 
we have presumed (in absence of any substantial evidence) that 
atrodorsalis does not cross it either. If this is the case, it may well 
be that the squirrel species Anderson mentions occurring in Marta- 
ban is in fact hyperythrus. He considered the type of hyperythrus to 
represent the phase of atrodorsalis without the black band on the 
back. We observed nothing about the type specimen of hyperythrus 
which mitigates against this concept (see description above), but it 
seems quite possible that squirrels taken just across the river from 
Moulmein at Martaban might in those days have been quite freely 
labeled "Moulmein," and that such material may in part constitute 
the phase of atrodorsalis without the black back which Anderson 
(1879, p. 233) describes from the Moulmein region. 

This problem will, of course, not be solved by disposition here of 
the name hyperythrus. What is needed is a series of specimens from 
Martaban and any points north of there (and west of the Salween) 
for comparison with the type of hyperythrus and with new material 
of the scarcer color phase of atrodorsalis (if any) from the Moulmein 
area and eastward from there toward Kawkereik and Myawadi. 

Callosciurus flavimanus siamensis (Gray) 

Sdurus siamensis Gray, 1860, Ann. Mag. Nat. Hist. (ser. 3), 5, p. 500. 
Sciurus atrodorsalis tachin Kloss, 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 178. 

Types. — Sciurus siamensis, BM No., young adult with 
dp* present, from Siam, collected by M. Mouhot; tachin, USNM 
No. 221566, a young adult female from Pak Bu, Tachin, Siam, taken 
on October 23, 1916, by C. Boden Kloss. 

Material examined, all from Thailand. — Bangkok (AMNH), three, 
(CNHM), six, (MCZ), 12, (UMMZ), 10, (USNM), 22; Pak Jong 
(AMNH), one; "Doi Souket," 6500 feet (MCZ), one; Paknampoh 
(MCZ), one; Tachin (USNM), six; Rajaburi (USNM), three; Pota- 
ram, near Ratburi (USNM), one; Ban Pong, near Rajaburi (USNM), 


Material examined, of intergrades with C. f. thai, all from Thai- 
land. — Paknampo, Nakorn Sawan (USNM), four; Moung Wat Sy, 
Nakorn Sawan (USNM), one; Kowkat, Paknampo, Nakorn Sawan 
(USNM), four; Kowkob, Paknampo, Nakorn Sawan (USNM), one. 

Material examined, of intergrades with C. f. pranis. — Tra Kha- 
nun, Kanachanaburi, Thailand (USNM), three; Hinlaem, Tra 
Khanun, Kanachanaburi, Thailand (USNM), one. 

The type locality of tachin seems to be the southwest edge of the 
geographic range of this subspecies, and material from there shows 
intergradation with pranis. Consequently, material from Tachin, 
Ratburi (=Rajaburi), Potaram, and Ban Pong are to be expected to 
show evidences of such intergradation. 

Diagnosis. — This subspecies is best known from the large series 
from Bangkok in several museums. C. /. siamensis is distinguished 
from C. f. thai on its west by having: (1) a maximum of four blackish 
bands on the tail hairs instead of five; (2) the tail increasingly red 
toward the tip instead of ivory or yellowish white; (3) the feet no 
blacker agouti than the legs; (4) the venter duller red, generally 
about Tawny or Hazel (although we recorded extreme ones as bril- 
liant as Burnt Sienna), whereas ventral pelage in six out of seven thai 
is Mahogany Red and the other Burnt Sienna; (5) smaller size; 
(6) no indication of a broad black longitudinal stripe on the poste- 
rior two-thirds of its mid-dorsum. 

From C. f. zimmeensis to the north, siamensis is also distinguished 
by above characters numbers 1, 2, 5, and 6. 

From C. f. pranis to the south, siamensis is distinguished by 
above characters numbers 2, 3, and 6. 

Callosciurus flavimanus pranis (Kloss) 

Sciurus erythraeus pranis, Kloss 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 178. 

Type.— USNM No. 221568, an old male from Koh Lak [Prachuap 
Kiri Khan], southwest Siam [Thailand], taken November 9, 1916, 
by C. Boden Kloss. 

Material examined, all from Thailand. — Koh Lak [Prachuap Kiri 
Khan] southwestern Siam (USNM), six, (NR), four, (BM), two; 
Hue Sai (NR), one; Sungei Balik (BM), one; Bok Pyin (USNM), 
two; Khao Luang, 3400 feet (ANSP), two; "Khao Nok Wua," 2000 
feet (ANSP), one. 

Pelage color. — The dorsal pelage is agouti and as a whole appears 
to be about Olive Brown and indistinguishable from that of tachin 


except that pranis regularly has an incipient expression of the black 
middorsal stripe that demonstrates its relationship to atrodorsalis. 
The reddish ventral pelage is regularly divided longitudinally by a 
narrow band of agouti pelage. The color of the reddish pelage ranges 
from Tawny Olive (XXIX) to Tawny. It also shows relationship to 
atrodorsalis and tachin in having the pelage of the rostrum hazel, 
and the ears tinged with the same color. 

Discussion. — This as yet poorly known subspecies is intermediate 
in some characteristics between C. /. atrodorsalis and rubeculus, but 
the material is difficult to shrug off as series of intergrades, because 
it seems consistently to possess a lighter, less richly colored venter 
than either atrodorsalis or rubeculus. 

Callosciurus flavimanus rubeculus (Miller) [extraterritorial] 

Sciurus rubeculus Miller, 1903, Smithsonian Misc. Coll., 45, p. 22. 
Sciurus erythraeus youngi Robinson and Kloss, 1914, Ann. Mag. Nat. Hist, 
(ser. 8), 13, p. 224. 

Types.— S. rubeculus, USNM No. 86777, old male from Khao Soi 
Dao, 1000 feet, Trong, collected February 21, 1899, by W. L. Abbott; 
youngi, BM No., old female, from Teku Plateau, 5000- 
6000 feet, Gunong Tahan, north Pahang, collected July 19, 1911, by 
H. C. Robinson and C. B. Kloss. 

Discussion. — Here is another subspecies which takes its name 
from a type specimen obtained in an area of intergradation with an 
adjacent subspecies. The typical material from Trong shows some 
trend toward intergradation with the only adjacent subspecies, pranis. 

Although C. f. rubeculus is partially isolated on a long peninsula 
far removed from most of the range of its species, it displays a most 
intensive expression of those conservative pelage characters which 
are so widespread in the species as to be regarded "typical" of the 
species. (No known pelage or other character has been found which 
delimits the species flavimanus.) These widespread characters are: 
faintly orange-rimmed ears, agouti feet blacker than the legs and 
sides, dark red ventral pelage bisected longitudinally by a narrow 
band of agouti gray pelage, tail hairs with four or five blackish bands. 

Diagnosis. — This subspecies is here distinguished from the only 
other conspecific one with which it is in contact, C. f. pranis, by: 
(1) absence of any suggestion of a broad blackish band on the poste- 
rior two-thirds of the mid-dorsum; (2) darker, richer ventral pelage; 
(3) blacker feet; (4) gray rostrum. 

Subspecies rubeculus is not mapped here. 


Callosciurus flavimanus hendeei (Osgood) 

Callosciurus erythraens hendeei Osgood, 1932, Field Mus. Publ. Zool., 18, 
p. 270. 

Type.— CNUM No. 32290, an adult male taken at Chapa, Ton- 
kin, February 14, 1929, by Russell W. Hendee. 

Material examined, from Tonkin. — Chapa, 5000 feet (CNHM), 
six, (MCZ), two, (USNM), one, (BM), 12; Bao Ha, 500 feet 
(CNHM), one, (BM), six; Muong Mo, 750 feet (CNHM), four, 
(USNM), one; Muong Boum (CNHM), one; Lieng San [Leng Sang], 
1500 meters (CNHM), one; Pakha [Pa Kha] (CNHM), two; Isle de 
la Table (BM), one; Lao Kay (MCZ), three; Thai Nien, 300 feet 
(BM), four; Lai Chau (USNM), one, (MNHN), one; Langson, 500 
feet (BM), one; Tam-dao (MNHN), one, (BM), four; Bac Kan 
(BM), four; "Chora," 1000 feet (BM), one; Ha-Giang (MNHN), 
one; Ngan-son, 3000 feet (MNHN), one; Muong Moun (AMNH), 

Material examined, from Laos. — Lao Fou Tchay, 3400 feet 
(CNHM). one; Phong Saly, 4400 feet (CNHM), four, (USNM), one; 
Muong Yo, 2300 feet (CNHM), two; Lo Tiao (MCZ), one; 2 km. 
east of Nam Khueng (MCZ), one; Hou^ Sai (MCZ), one. 

Material examined, from Annam. — Phu Qui, 100 feet (CNHM), 
two, (MNHN), one, (BM), seven; Than Hoa (CNHM), one; Hoi 
Xuan (CNHM), one; "Lung-lunh" (CNHM), two, (BM), one; Lung 
Van, 1000 meters, pr. de Thanhoa (USNM), one; "Nam Da, near 
Tatka" (BM), one; Nghia Hung, 100 feet, Phu Qui (BM), seven. 

Material examined, from China. — "San-ho Hsien," Kweichow 
(CNHM), one; "Shui-kow-kwan," Lungkow, Kwangsi (CNHM), one. 

For discussion of characters and relationships see castaneoventris 
and flavimanus. 

Callosciurus flavimanus castaneoventris (Gray) 

Sciurus castaneoventris Gray, 1842, Ann. Mag. Nat. Hist. (ser. 1), 10, p. 263. 
Sciurus erythraeus insularis J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 
22, p. 473. 

Types. — Sciurus castaneoventris, BM No. 72A, young female and 
72B adult male, both from "China," evidently the vicinity of Canton 
and collected between 1812 and 1831, for they were presented by 
John R. Reeves (who worked there as a tea taster during that time; 
see Moore and Tate, 1959) ; S. e. insularis, AMNH No. 26609, parous 
female from "Lei-Mui-Mon," Hainan, taken January 5, 1903 by 
agents of Alan Owston. 


Material examined, all from Hainan, China. — "Lei Mui Mon" 
(AMNH), 14; "Utocki" (AMNH), one; "Luidon" (AMNH), one; 
"Hainan" (AMNH), three; Nodoa (AMNH), 80, (USNM), two, 
(CNHM), 14, (BM), one, (MCZ), six; Nam Fong (AMNH), seven, 
(CNHM), two; Kachek (USNM), one; "Ngau Tchi Lea Mts." 
(MCZ), one, (BM), one. 

Discussion. — No specimens of the species flavimanus, other than 
the cotypes collected by Reeves are known from the vicinity of Can- 
ton, but Robinson and Kloss (1918, p. 199) compared the cotypes 
with material from Hainan and found them to be indistinguishable, 
thus rendering J. A. Allen's subspecies insularis from Hainan a syn- 
onym of castaneoventris. Without comment these authors thereupon 
substituted Hainan as type locality for the "China" which was the 
only type locality provided by Gray in his original description of 
castaneoventris. While that substitution was a considerable improve- 
ment in its time, as indicated above, it now seems better to accept 
the vicinity of Canton as type locality, but only provisionally until 
further material becomes known from there. 

There were two specimens involved in the original description of 
castaneoventris. The first mentioned was a rather darker one (BM 
No. 72B) the original description of which amounts to, "Very like 
S. hippurus, but only half the size, and the ears are gray." This im- 
plies red ventral pelage like that of hippurus, for Gray proceeds to 
contrast the variant second specimen with it, "Var. rather paler; 
chin grayish, beneath yellowish red." The amount of variation im- 
plied does not require that the type series came from widely different 
localities nor that it came from a single locality which is in an area 
of intergradation between two subspecies (castaneoventris and ning- 
poensis) . Nevertheless, the location of Canton, from which the type 
material is presumed to come, in reference to known localities for 
ningpoensis and subsequent castaneoventris material does suggest this 
latter possibility, and the Hungshui River which debouches about 
Canton and Macao, could quite possibly form a partial, natural bar- 
rier between the two subspecies. If this should prove to be the case, 
the type series does better represent the southern, not the northern 
subspecies, and no need is anticipated to alter the well-established 
usage of castaneoventris for Hainan material and ningpoensis for that 
from Fukien. 

In his diagnosis of the subspecies hendeei Osgood (1932, p. 270) 
stressed no single character which could be depended upon alone to 
distinguish it from the one on Hainan, but stated all the differences 


in subjectively comparative terms. It seems worthwhile to provide 
quantitative differences here as we find them to exist. 

Pelage color. — Among the large series of castaneoventris from Hai- 
nan at the American Museum of Natural History there are only five 
individuals that have a full-length midventral band of agouti pelage. 
There are 56 that have agouti throats and an agouti wedge extending 
onto the breast, and 41 that have neither midventral band nor wedge. 
This contrasts with the series of thirty skins of hendeei at Chicago 
Natural History Museum in which there were no full-length, ventral, 
median bands of agouti and only one with an agouti wedge on the 
breast (No. 32297). 

The ventral pelage color of the large AMNH series from Hainan 
is about Mahogany Red in about 30 specimens. There are one or 
two skins of adults in good pelage as pale as Mars Orange, and the 
remainder are about Burnt Sienna (except for one individual from 
Nodoa (AMNH No. 58133) which has such an exceptionally agouti 
venter that virtually only the insides of the legs have really red pel- 
age. The possibility of such aberrant individuals elsewhere needs to 
be borne in mind in this species) . This is a very high degree of con- 
sistency in ventral pelage color compared with ningpoensis. In the 
(smaller) Chicago collection of hendeei material, however, the ven- 
tral pelage color was found to vary from about Morocco Red to Hays 
Russet, a range of variation which is even more conservative than 
the castaneoventris. 

C. f. castaneoventris and C. f. hendeei both have five black bands 
on the tail hairs. 

The most constant and obvious color difference between the hen- 
deei and castaneoventris as represented in the large series at Chicago 
and New York is the color of the tip of the tail. In the Hainan ma- 
terial there appears to be no variation away from a white-upon-black- 
tipped tail. Some description is necessary here. The tail hairs of 
these C. f. castaneoventris from Hainan, excepting near the distal end 
of the tail, have five blackish bands and a minute blackish tip as is 
widespread in the species. But the light subterminal band is un- 
usually light in this species (about Cartridge Buff) and is progres- 
sively longer toward the extremity of the tail, often exceeding a 
centimeter at the extremity, but rarely two centimeters. The most 
distal black band becomes progressively longer, also, toward the end 
of the tail, by spread toward the base of the hair. Thus, at the 
end of the tail the subterminal whitish band of one centimeter is 
distal to a sub-subterminal black band of from two to five centi- 


meters. (Osgood, incidentally, ignored the minute blackish tips of 
the hairs and described the whitish bands as terminal.) 

The tips of the tails of the hendeei series at Chicago Natural 
History Museum are characteristically and uncomplicatedly black. 
There are four (three from Chapa and one from Lung-lunh), how- 
ever, which do have long, light bands superimposed on the extrem- 
ities of the black hairs, rendering them indistinguishable by this 
criterion from Hainan material. However, 87 percent is a fairly good 
distinction of subspecific material (Amadon, 1949) on the basis of a 
single color character. 

Dimensions. — Size is something on which we can also provide 
data: Greatest skull length in 36 adults of castaneoventris ranges 
from 50.2 to 53.5 mm., except for two well outside the curve at 55.0 
and 55.9, and 29 (80 percent) are less than 53 mm. long; whereas all 
17 skulls of adult hendeei are 53 mm. or more in length. Similarly, 
the hind foot length of the adults of hendeei ranges from 54 to 59 mm., 
but in a sample of 61 adult castaneoventris only four range above 
53 mm. 

Callosciurus flavimanus flavimanus (Geoffroy St. Hilaire) 

Sciurus flavimanus Geoffroy St. Hilaire, 1831, in Belanger, Voyage Indies 

Orientales, 1, p. 148. 
Callosciurus flavimanus quantulus Thomas, 1927, Proc. Zool. Soc. London, 

1927, p. 51. 
Callosciurus flavimanus contumMX Thomas, 1927, Proc. Zool. Soc. London, 

1927, p. 52. 
Callosciurus flavimanus dactylimis Thomas, 1927, Proc. Zool. Soc. London, 

1927, p. 52. 
Callosciurus flavimanus piraia Thomas, 1929, Proc. Zool. Soc. London, 1928, 

p. 836. 
Callosciurus flavimanus bolovensis Osgood, 1932, Field Mus. Nat. Hist., Zool. 

Ser., 18, p. 276. 

Types. — S. flavimanus, MNHN No. 177, from Tourane, Annam; 
quantulus, BM No., adult male from Xieng Quang Koo, 
Laos, collected December 19, 1925, by J. Delacour and W. Lowe; 
contumax, BM No., adult female from Kontoum, An- 
nam, collected February 27, 1926, by J. Delacour and W. Lowe; 
dactylinus, BM No., old female from Dak-to, Annam, 
collected March 16, 1926, by J. Delacour and W. Lowe; pirata, BM 
No., adult female from Nape, 2500 feet, Laos, collected 
February 11, 1928 by J. Delacour and W. Lowe; bolovensis, CNHM 
No. 37874, old female from Paksong, Indochina, collected Janu- 
ary 24, 1932 by J. Delacour. 


Material examined, from Annam. — Col de Nuages, 400 meters 
(MNHN), eight, (BM), 12; Thua Luu, 150 feet (BM), two; Dak To 
(BM), two, (MNHN), two; Kao Hao, Quangtri Province (BM), one; 
Kontoum [Kontum] (BM), two, (MNHN), two; Hu^ (BM), one; 
Cua-Ting, Quangtri (BM), one; Quangtri (BM), one, (MNHN), 
one, (CNHM), six, (USNM), one; Dong Hoi (MNHN), one; Quang- 
tri River (MNHN), one. 

Material examined, from Laos. — Thateng (CNHM), 15, (BM) 
one, (USNM), one; Nap6, 2500 to 3000 feet (BM), five; Xien Kuang 
Koo [Xien Khouang] (BM), three, (MCZ), two; Pasa (AMNH), one; 
Don Qua (AMNH), one; Plateau Bolovens (AMNH), four; Phu 
Kobo (MCZ), two; Bantion [Ban Tayun] (CNHM) one; "Pakhout" 
(CNHM), one; Pak Song (CNHM), one; Nong Kai [presumed to 
mean in Laos across the river from Nong Kai, Thailand] (USNM), 

Discussion. — The two AMNH specimens from Pasa and Don 
Qua, villages on the Nam Khan (river) between Luang Prabang and 
M. You, exhibit a condition of intermediacy between C. f. hendeei 
and C. f. flavimanus. There is no geographic barrier between these 
localities and those of the hendeei. There is no difference from hen- 
deei in the appearance or size of the Pasa and Don Qua skulls, and 
body measurements of these and hendeei taken by the same collector 
(T. Donald Carter) offer no hint of a size difference. The rostrum 
is colored like the dorsum as in hendeei rather than orange-yellow 
as in flavimanus. The feet are agouti like the back, a stage midway 
between the blackened agouti of hendeei and the orange-yellow of 
flavimanus. However, the Pasa specimen possesses pronounced black- 
ish agouti on its toes showing more influence of the hendeei character; 
whereas the Don Qua one lacks the blackish agouti entirely but has 
a faint infusion of reddish-orange around the ankles (absent in the 
Pasa specimen) showing more the influence of flavimanus. In both 
of these Nam Khan (river) specimens the tail is colored like the back 
but brighter buff approaching the condition of flavimanus, not black- 
tipped like hendeei. 

The Museum of Comparative Zoology material from Xien Khou- 
ang and Phu Kobo seems to be quite good flavimanus with no indi- 
cation of intergradation ; but five of the six Chicago Natural History 
Museum specimens from Quangtri have abrupt solid black tail tips 
indicating some penetration of hendeei characteristics even to that 
locality. Both of the Nam Khan specimens have four prominent 
blackish bands on the tail hairs. This is a reduction of one from the 


number characteristic of hendeei and one or two more than is com- 
mon in flavimanus farther south. 

The magnitude of difference between diagnostic characteristics 
of flavimanus and hendeei does greatly exceed that between hendeei 
and any of the Chinese subspecies. The evidence of intergradation 
described above compells us, however, at this state of our knowledge, 
to regard subspecies flavimanus and hendeei as conspecific. It is also 
quite evident to us that flavimanus material is much more variable 
individually than is that of hendeei, but these conditions in no way 
justify admission as good subspecies any of the many other names 
that have been applied in this case. 

Pelage color. — The characteristics accepted for flavimanus are 
orange-yellow rostrum and fore feet, and reddish-orange hind feet. 
The tail may be colored like the dorsum or huffier or approaching 
the color of the venter. The venter varies among individuals from 
Mars Orange to Burnt Sienna. This color extends from wrist to 
ankle and covers the venter except for the agouti scrotum and chin. 
An occasional individual shows feeble anterior indications of a mid- 
ventral agouti stripe. Faint indications of a broad, black midstripe 
on the posterior two-thirds of the dorsum is characteristic. 

Callosciurus flavimanus griseimanus (Milne-Edwards) 

Sciurus griseimanus Milne-Edwards, 1867, Rev. Zool. (ser. 2), 19, p. 195. 
Macroxus leucopus Gray, 1867, Ann. Mag. Nat. Hist. (ser. 3), 20, p. 282. 
Sciurus leucopus fumigatus Bonhote, 1907, Abstr. Proc. Zool. Soc. London, 

no. 38, p. 2. 
Sciurus vassali Bonhote, 1907, Proc. Zool. Soc. London, 1907, p. 9 (footnote, 

a new name for fumigatus Bonhote, 1907, not of Gray, 1867). 
Callosciurus ferrugineus phanrangis, Robinson and Kloss, 1922, Ann. Mag. 

Nat. Hist. (ser. 9), 9, p. 91. 

Types. — Sciurus griseimanus, MNHN No. 1864-679(167), adult 
male from Saigon, collected by R. Germain; leucopus, BM No., adult male from Cambodia; vassali, BM No., 
male, from Bali, near Nhatrang, Annam; phanrangis, BM No. 26.- 
11.17.5, adult female from Tour Cham near Phanrang, south Annam, 
collected May 23, 1918, by C. B. Kloss. 

Material examined, from Annam. — Djiring, 3000 feet (BM), 11, 
(MNHN), four; Da Ban (BM), one; Nhatrang (BM), 12; Phanrang 
(BM), one; Lang Bian Peaks (MCZ), one; Pie de Langbian (MCZ), 
one; forest of Krongba [Krong Ba] (USNM), one. 


Material examined, from Cochin China. — Tayninh (BM), four, 
(MNHN), six; An Binh (BM), three; Trang Bom (BM), two; An 
Blu (MNHN), one; "Arboretum de" Trang Bom, Province de Bien 
Hoa (USNM), three; Bien Hoa (USNM), five; (MNHN), one; 
"Montaojur de Nrii," Chua Chan (USNM), one; "Prang," (USNM), 
one; Baria (CNHM), two; Ninh Hoa (CNHM), two; Phan Rang 
(CNHM), two; "Gougah" (CNHM), two; Ban Me Thuot (CNHM), 
two; "Eaktur" (CNHM), one. 

Diagnosis. — The distinguishing characteristics of griseimanus are: 
(1) very Hght dorsal pelage, some Mouse Gray, some Drab; (2) seem- 
ingly white feet, about Cream Buff in some. Cartridge Buff in others; 
(3) tails not tipped with a contrasting color, but unusually disposed 
to show transverse annulations, numbering about ten; (4) number 
of blackish bands on tail hairs at least four, often five. 

There is, in griseimanus, as in flavimanus, a great deal of varia- 
tion that is individual or microgeographic or both. 

Pelage color. — The ventral pelage color in eleven specimens at the 
CNHM varies from Sanford's Brown in two, through Ochraceous 
Buff in two. Cinnamon Buff in five, to something lighter still in two 
immatures from Phan Rang. Bisection of the ventral pelage by an 
agouti stripe is incipient in four of these eleven. 

Discussion. — A specimen (USNM 256749) indistinguishable from 
hendeei was, according to the label, taken by Poilane on Nua Chua 
Chan, a small mountain close to Bien-hoa, Cochin China, deep in 
the range of subspecies griseimanus, from which hendeei differs 
greatly. A companion specimen of hendeei was taken by Poilane 
(USNM 256748) at Lung Van, Than Hoa Province, Annam, well 
within the proper range of hendeei. It seems rather more likely that 
the first specimen somehow became erroneously labeled than that a 
squirrel which is indistinguishable from hendeei could in fact have 
been taken on Nua Chua Chan (see map of species flavimanus, figure 

The color difference between subspecies griseimanus and flavi- 
manus is just as striking as that between subspecies flavimanus and 
hendeei, but it is color intensity which differs more than pattern, and 
in view of the color patterns common to these first two forms and 
absence of any geographical barrier between them, it would seem 
unrealistic to us even in the absence of noticed intergradation, to 
regard griseimanus and flavimanus as more distinct than subspecies. 


Callosciurus flavimanus gloveri (Thomas) 

Callosciurus castaneoventris gloveri Thomas, 1921, Jour. Bombay Nat. Hist. 
Soc, 27, p. 502. 

Type.— Callosciurus castaneoventris gloveri, BM No., 
young female, from Nagchuka, 10,000 feet, west Szechwan, col- 
lected August 14, 1908, by W. R. Zappey. 

Material examined, from Szechwan, China. — West of Fulin, 5000 
feet (USNM), one; Ningyuanfu, 6200 feet (USNM), one; "Tanken" 
(USNM), one; 40 miles E. of Hokow (ANSP), one; Hokow (=Nag- 
chucka) (ANSP), one topotype, Nagchucka, 10,000 feet (BM), one 
topotype; "Toloko," south of Muli (MCZ), one, (ANSP), one; Na- 
chukar [Nagchuka] (MCZ), three topotypes, (USNM), one; Rama 
La Pass, 13,000 feet (MCZ), three; Mili [same as Muli] (CNHM), 
three, (USNM), three; Baurong (CNHM), four, (ANSP), one. 

Material examined, from Yunnan, China. — Yungning (CNHM), 
two; Yung-pei-ting, "N. 27° 10', E. 100° 50'" [Yungpeh] (BM), one. 

Material examined, from Tibet. — "U-long-si Gorge," 14,000 feet 
(USNM), two. 

Pelage color. — Color notes from material examined in the Museum 
of Comparative Zoology and Chicago Natural History Museum that 
includes three topotypes: The ventral pelage varies from Hay's Rus- 
set (XIV) through Vinaceous-Rufous to Ferruginous from individ- 
ual to individual, but is consistent on any one specimen from throat 
to tail and wrists to ankles, and is not in any instance divided by a 
mid-saggital agouti band, even incipiently. Dorsal pelage an agouti 
of about Mouse Gray (LI) at the more northern localities of Nag- 
chuka and Rama La Pass, Grayish Olive (XLVI) at Baurong, and 
Dark Olive Buff (XL) at the more southern localities of Mili and 
Yungning. Inner side of pinna is Apricot Buff (XIV) in the more 
northern material and Cinnamon Rufous to Vinaceous Rufous in 
the southern, but at Baurong there is just a hint of warm color in the 
gray. The pelage of the feet is colored like that of the dorsum. The 
tail is colored like the dorsum for its full length, and the tail hairs (at 
Yungning, Mili, and Baurong, at least) have six dark bands, and tips 
of Ochraceous Buff (at least in the north) . 

Discussion. — This rather variable subspecies shows relationship to 
intermedius to the west and bonhotei to the east in having undivided 
ventral pelage and uniformity in color of tail. It differs from both 
of them in having reddish ears and unblackened feet. Its dorsal 
pelage is also paler than that of intermedius and lacks the reddish 



Callosciurus flavimanus bonhotei (Robinson and Wroughton) 

Sciurus castaneiventris bonhotei Robinson and Wroughton, 1911, Jour. Fed- 
erated Malay States Mus., 4, p. 234. 

Type.— BM No., an adult female taken at Chin Chien 
San, Szechwan, China, by F. W. Styan. 

Material examined, all from Szechwan, China. — Lian-feng-Kiang, 
Omei Shan (CNHM), one; Yaichowfu [Yachowfu], Omei Shan 
(CNHM), two, (USNM), one, (ANSP), three; "Ho Ni Pa" (CNHM), 
nine; Fi Shan Kwan (CNHM), eight; Lu Chang Pu (CNHM), two; 
Yung Cha Shan (CNHM), one; Chung Chiao [Chiang] Miao (CNHM), 
one; Hsiao Yang Chi (CNHM), one; Fu Fu Su (CNHM), three; 
Ninguanfu (USNM), one, (BM), one; "Tai Tsin Tang" (USNM), 
one; "Wan Nien Si" (USNM), one; "Lu Din" (USNM), one; Kia- 
ting (USNM), two; Mt. Omei (USNM), four; Suifu (USNM), two; 
"Shin Kai Si," Mt. Omei (USNM), five; Wan hsien (MCZ), one; 
"Chen Yen Say" (BM), one; "Yusu-Ching-Hsien" (BM), one; 
"Szechwan" (BM), two. 

There is a specimen referable to gloveri as well as one referable to 
bonhotei, both taken at Ningyuanfu and both showing some indica- 
tions of intergradation. As the map shows, there is one other point 
at which individuals bearing the characteristics of these two sub- 
species were taken very close together. 

Pelage color. — The agouti dorsal pelage of the bonhotei material 
at Chicago Natural History Museum is generally about Dresden 
Brown (XV) with the reddest variant being Sudan Brown (III). 
The tail hairs have six dark bands and sometimes seven. The ven- 
tral pelage is Burnt Sienna and is not divided in any of this series. 
Very little warm color can be found on the ears. 

Discussion. — Although somewhat variable, the material reported 
above appears to represent a distinct enough geographic subspecies 
which is richer in ventral pelage and darker in dorsal pelage than the 
adjacent gloveri, and also differs by having its feet slightly blackened 
and the reddish orange color almost absent from its ears. Resem- 
blance to hendeei of eight specimens from Fi Shan Kwan and two 
from Lu Chang Pu in characters of dorsal pelage is noted. However, 
these are the most northwestern and most southeastern of the col- 
lecting localities found. In the Lu Chang Pu two the tail has a long, 
black subterminal mark with the tip itself buffy or whitish, which 
is also like the northern hendeei known to us. In the other 19 ex- 
amples of bonhotei in the Chicago Natural History Museum series, 


however, difference from hendeei in dorsal pelage characters is fairly 

It is interesting to note that the northern limit suggested by the 
collecting localities mapped here may be close to correct. The Sage 
West China expedition of the American Museum of Natural History 
collected west and north out of Wenchwan for a total of about two 
months without taking squirrels of this species. They obtained 
series of other squirrels (Dremomys, Sciurotamias, and Tamiops) and 
good stands of coniferous forest and bamboo forest in the vicinity 
of their camps are evident in photographs shown to us by T. Donald 
Carter of that expedition. Most of their collecting was above 6000 

Callosciurus flavimanus styani (Thomas) 

Sciurus styani Thomas, 1894, Ann. Mag. Nat. Hist. (ser. 6), 13, p. 363. 

Macroxus griseopectus Milne-Edwards, 1874, Recherches pour servir a I'His- 
toire Naturelle des Mammiferes ... p. 305 (not griseopectus Blyth, 1847). 

Herpestes leucurus Hilzheimer, 1905, Zool. Anz., 29, p. 299. 

Herpestes albifer Hilzheimer, 1906, Abh. Ber. Mus. Nat. Heimatk. Magde- 
burg, 1, p. 177 (substitute for H. leucurus, preoccupied). 

Callosciurus caniceps canigenus Howell, 1927, Jour. Washington Acad. Sci., 
17, p. 81. 

Callosciurus erythraeus woodi Harris, 1931, Occas. Papers Mus. Zool., Univ. 
Michigan, no. 228, p. 1. 

Types, all from China. — S. styani, BM No., young 
female "from between Shanghai and Hangchow [about 100 miles], 
probably Kahing ..." [Kashing]; griseopectus, collected by A. David 
in Chekiang; leucurus, "2 Felle."; albifer, "2 Felle. In Hankau 
gefauft. 1905"; C. canigenus, USNM No. 241509 an adult male 
taken at Hayenhsien, Hangchow Bay, Chekiang, December 10, 
1925, by Arthur de C. Sowerby; woodi, UMMZ No. 55827, adult 
male, taken at Lungtan in the Purple Mountains 25 miles east of 
Nanking, Kiangsu, south side of Yangtse River, December 14, 1923, 
by Norman A. Wood. 

Material examined, all from China. — Tehan (UMMZ), one; Wan 
Mts., 100 mi. S.W. of Nanking, Anhwei (UMMZ), five, (MCZ), two; 
Purple Mts., Lungtan, 25 mi. E. of Nanking, Kiangsu (UMMZ), 
three; Tunglu, Chekiang (AMNH), one, (MCZ), nine; Mokanshan, 
Chekiang (ANSP), three; Hai Yen Hsien, Hangchow Bay, Chekiang 
(USNM), two; Kangpu, near Hangchow, Chekiang (USNM), one; 
Nimrod Sound, coast of Chekiang (ZMHU), one; "Anhwei" (NR), 
17; "Chien San," Anhwei (BM), three; Lushan Hills, Kiukiang, 


Kiangsi (BM), one; "Hung Fa Chao," Shanghai (UMMZ), one; 
"Kin King," Chekiang (USNM), one. 

We are in agreement with G. M. Allen (1940, p. 633) on the above 
synonymy, which he has fairly and adequately discussed (pp. 634, 

Pelage color. — The series from the Wan Mountains are dorsally 
an agouti of Light Brownish Olive (XXX) in most, but one is Buck- 
thorn Brown (XV) . There are five blackish bands on the individual 
tail hairs. There is an abrupt, short tip of black hairs present on the 
tail, but this is obscured to some extent by whitish or buffy tips even 
on the otherwise entirely black hairs. No orange is seen on the ears 
nor black on the feet. The ventral pelage is Light Ochraceous Buff 
to Pale Ochraceous Buff. Chin or chin and throat are agouti like 
the dorsum. There are two from the Purple Mountains with ven- 
tral pelage that is Pale Pinkish Buff to almost White, and their dorsal 
pelage is agouti of about Dark Olive Buff (XL) which is very slightly 
lighter than the lightest of the series from the Wan Mountains. The 
other Purple Mountains specimen, however, has blackish feet, dorsal 
pelage agouti of about Dresden Brown, and ventral pelage of Mo- 
rocco Red. All three from the Purple Mountains have chin or chin 
and throat agouti and ears concolorous with dorsum. 

Discussion. — Thomas (1894) distinguished styani from "castaneo- 
ventris" to the south by the ventral pelage being a "peculiar reddish 
cream-colour ('pinkish-buff' of Ridgway) instead of the rich rufous 
('orange-rufous') of the older known form." A good deal more mate- 
rial has since become available, and although some variation toward 
ningpoensis characteristics is seen in styani, the difference is still as 
Thomas said "extremely striking." When the material is compared 
in series, a general difference is also evident in the color of the dorsal 
pelage, but there is some overlap among individuals. As Thomas 
said also, in other pelage characters the two subspecies seem to be 
quite alike. 

Harris (1931) proposed the name woodi for material from east of 
Nanking in the Purple Mountains which has lighter pelage in both 
dorsum and venter than previously found in styani. However, some 
years later he obtained another specimen (UMMZ No. 87081) from 
the same vicinity with dorsal pelage about Dresden Brown (XV) 
and ventral pelage that is about Morocco Red (I). This must be 
regarded as a most exceptional specimen for styani, and it certainly 
invalidates woodi as a subspecies. 


Since the subspecies styani is known from quite close to the south 
of the Yangtze Kiang in several places but as yet from no localities 
north of that great river, it seems possible that the lower course of 
the Yangtze in eastern China has been a barrier to northward spread 
of C. flavimaniLs. 

Callosciurus flavimanus ningpoensis (Bonhote) 

Sciurus castaneoventris ningpoensis Bonhote, 1901, Ann. Mag. Nat. Hist, 
(ser. 7), 7, p. 163. 

Sciurus tsingtanensis Hilzheimer, 1905, Zool. Anz., 29, p. 298 ("corrected" to 
tsingtauensis, 1906, Abhandl. Berichte Mus. Natur. Heimatk. Magde- 
burg, 1, p. 172). 

Types. — Sciurus c. ningpoensis, BM No., old individ- 
ual from hills 30 miles from Ningpo, collected March, 1884, by F. W. 
Styan; tsingtanensis, ZMHU (Kreyenberg) No. 238, a male collected 
August 16, 1902, by Kreyenberg (at Nimrod Sound, coast of Che- 
kiang, according to G. M. Allen, 1940, p. 632) just south of Shanghai 
(Tsingtao is 400 miles farther north). 

Material examined, all from China. — Ningpo, Chekiang (AMNH), 
eight, (BM), seven, (MCZ), two, (CNHM), two; Fuching Hsien 
[Futsingl, Fukien (AMNH), 53, (MCZ), six; (CNHM), four; Yen- 
ping, Fukien (AMNH), six, (CNHM), two; Chungan Hsien, Fukien 
(AMNH), four; Kushan [Kaoshanshih] near Foochow, Fukien 
(USNM), one; "Peiliang," near Foochow, Fukien (USNM), one; 
Foochow [later Minhowl (CNHM), one, (BM), six; "Fukien" (NR), 
one, (CNHM), six; "Ah Chiung," Fukien (BM), three; "Mayhos," 
China (BM), one; Pootoo I. [Putu Shan], Chusan Archipelago, Che- 
kiang (BM), one. 

Pelage color. — The series of eight topotypes of ningpoensis in the 
American Museum of Natural History is consistently lighter colored 
in dorsal pelage than the majority of the 63 specimens from several 
localities in Fukien Province. While the range in ventral pelage 
color is very similar between the Ningpo and the Fukien material, 
the frequency of the palest color, about Apricot Buff, in the Ningpo 
is 50 per cent and in the Fukien three per cent. The richest ventral 
pelage color in the Ningpo eight is about Ferruginous, occurring in 
three specimens. Forty-one per cent of Fukien AMNH specimens 
have a richer pelage than Ferruginous, the richest color attained be- 
ing about Sanford's Brown. 

In 16 per cent of the AMNH Fukien series there is a longitudinal 
midventral band of gray agouti pelage entirely bisecting the red pel- 


age. In 56 per cent there is an agouti wedge on the throat and breast 
dividing only the anterior portion of the red pelage. In 30 per cent 
there is no division of the red pelage. Of the eight Ningpo specimens 
five have an agouti wedge and three no division at all. The ventral 
pelage of the Tunglu series of nine at the Museum of Comparative 
Zoology is generally about Sanford's Brown with one about Orange 
Cinnamon (XXIX) and another Ochraceous Buff (XV), although 
the AMNH Tunglu specimen is quite as pale as styani, Light Ochra- 
ceous Buff at its richest. 

It should be said that material included in this subspecies seems 
to have six black bands to the fully grown-out tail hairs, besides the 
hardly noticeable blackish tip. And black-tipped guard hairs with 
three (as well as two, one, and no) light bands, occur commonly in 
the dorsal pelage. 

Discussion. — A pale subterminal band on the tail hairs is so long 
and noticeable that it gives a strong impression that the tail hairs 
are pale-buff tipped. The terminal hairs of the tail are each black 
for most of their length proximal to the mentioned subterminal light 
band. This creates a sub-sub terminal black spot in the tail. This 
spot is particularly noticeable from the under side and quite like that 
found also characterizing gordoni and michianus. It rarely shows 
any disposition to spread onto the sides of the tail as is seen in hendeei 
and castaneoventris. This black tail spot thus suggests closer rela- 
tionship of ningpoensis to michianus (and through it to zimmeensis 
and gordoni) than to the subspecies honhotei and gloveri north of the 
Yangtze Kiang. The above evidence of relationship finds striking 
support in the common possession by ningpoensis, michianus, zim- 
meensis, and gordoni, of the agouti wedge on the breast or a full- 
length mid ventral agouti band. North of the ranges of these four 
subspecies, gloveri and honhotei each have undivided red ventral pel- 
age and uniformly colored tail. To the south of the same four sub- 
species, the agouti wedge on the breast (as well as the full-length 
midventral agouti band) becomes scarce in castaneoventris and ab- 
sent in hendeei. Correspondingly, the tails of these two subspecies 
are distinguished from the black spot characterizing ningpoensis, 
michianus, zimmeensis, and gordoni. In hendeei and castaneoventris 
black is not confined to a terminal spot but extends anteriorly along 
each outer edge of the tail, progressively diminishing anteriorly. 

Callosciurus flavimanus thaiwanensis (Bonhote) 

Sciurus thaiwanensis Bonhote, 1901, Ann. Mag. Nat. Hist., (ser. 7), 7, p. 165. 


Sciurus thaiwanensis centralis Bonhote, 1901, Ann. Mag. Nat. Hist., (ser. 7), 

7, p. 166. 
Sciurus thaiwanensis roberti Bonhote, 1901, Ann. Mag. Nat. Hist., (ser. 7), 

7, p. 166. 
Callosciuru^ erythraeus nigridorsalis Kuroda, 1935, Jour. Mammal., 16, p. 281. 

Types. — S. thaiwanensis, BM No., adult male from 
Baksa, Formosa, collected October 20, 1893, by P. A. Hoist; cen- 
tralis, BM No., adult female from Lak Ku Li, central 
Formosa, collected June 29, 1894, by P. A. Hoist; roberti, BM No., adult male from northwest Formosa, collected by Robert 
Swinhoe (skin only); nigridorsalis (not seen), Marquis Yamashina 
Collection No. 22, adult female from Riran, Taito, southeast For- 
mosa, collected August 1, 1932, by H. Orii. 

Material examined, all from Taiwan (Formosa) . — Teraso [Teraso 
Soan] (AMNH), two; Chip Chip (AMNH), three; Mt. Arisan, 8000 
feet (MCZ), one, (BM), four; Kagi District (CNHM), three; "For- 
mosa" (UMMZ), three; Tainan (UMMZ), one; Racu Racu [Raku- 
raku] Mountains, 7000 feet (BM), two; Tapposha [Tappan-sha], 
Central Formosa (BM), five; "Ho Ho Mountain," 5000 feet (BM), 
one; "Lak-ku-H," Central Formosa (BM), three; Baksa (BM), one; 
"Lau Long" (BM), one; South Cape of Formosa (BM), two; Nan- 
Tou Hsien, Wu-Sheh (AMNH), one; Ping Tung (AMNH), one; 
Taipei Hsien, Wu Lai, 16 miles south of Taipei (AMNH), five; 
"I-Lan Hsien," Chiao-chi (AMNH), two; Taipei (AMNH), one; 
"Taipei Hsien," Ping Ling, Taipei (AMNH), five; I-Lan Hsien, 
I-Lan, Taipei (AMNH), two. 

Discussion.— Bonhote (1901) described three Formosan forms (of 
flavimanus) from only three localities and apparently very few, if any, 
more than three specimens. Furthermore, the localities from which 
Marquis Nagamichi Kuroda (1935, pp. 280-282) reported specimens 
in support of Bonhote's concepts are exceedingly few: one for roberti, 
two for centralis, one for thaiwanensis, and two for the subspecies he 
himself proposed, nigridorsalis. Kuroda did have, and enumerate, 
at least modest series from these localities, and does to some extent 
describe the variation noted in his series. The material that we have 
been able to examine, however, does not support all of Kuroda's 

The only Baksa we find is about 95 miles north of the southern 
tip of Taiwan. We presume this Baksa to be the type locality of 
thaiwanensis. This type locality is farther north than the entire 
known range of nigridorsalis, although Kuroda says that thaiwanen- 


sis "is found only in the southernmost parts of the island" implying 
south of the range of his nigridorsalis. If Kuroda's implication is 
correct, thaiwanensis would appear to be restricted to the peninsula 
of the southern cape, failing by 40 or 50 miles to reach its own type 
locality. Kuroda (1935) did not establish whether thaiwanensis, in 
his restricted concept, may in fact extend from the south peninsula 
westward along the south coast and then northward toward Baksa. 
The specimen from Ping Tung, having a red venter, strongly sug- 
gests that the gray-bellied population does not so extend westward 
and northward. If the locality we find for Baksa is the correct one, 
the type is therefore but a variant specimen from well within the 
geographic range of the centralis population. This concept of the 
type is supported by the other Baksa specimen in the British Mu- 
seum which one of us (Tate) found to be like the centralis population. 
The above four specimens from Teraso and South Cape support 
Kuroda's finding that a population exists on the South Cape in which 
the venter is consistently (May, November, January) all gray or has 
but small patches of reddish at the bases of the limbs. 

The greatest known distance between two localities from which 
only the grey-bellied south cape population is known, is between the 
"South Cape" and Koshun, ten miles. The distance between the only 
two known localities for nigridorsalis, Taito and Daibusan, is thirty 
miles. Only one specific locality has been attributed by a taxonomist 
to the dorsally reddish subspecies roberti, that is, Taiheizan. This 
reddish infusion is quite evident in one male specimen from Taipei 
Hsien, Ping Ling, but absent from two other males taken there the 
same day. The reddish infusion is quite evident in two specimens 
from Taipei Hsien, Wu Lai, but not in four others from the same 
place. The alleged differences between all these subspecies may be 
geographically constant enough over a large enough range to merit 
recognition as respectable geographic subspecies, but it seems to us 
that the constancy in some cases and the range in all remains to be 

Diagnosis. — Bonhote (1901, p. 165) pointed out certain features 
that characterize all C. flavimanus on Formosa, and which may be 
altered to read: (1) Hairs of the dorsal pelage have 2 or 3 light bands. 
(2) The ears are somewhat orange on the concave surface. (3) The 
tail is agouti (with 3 to 5 blackish bands on each hair) for the prox- 
imal half-length but blackens progressively out the distal half, where 
it also has on each hair a long, yellowish tip. (4) In addition, the 
variation in ventral pelage is like that described above for castaneo- 


ventris except that the percentage of each kind of variant differs. 
In a sample of 23 skins four have red throats and no, or virtually no, 
agouti in the midline, eleven have agouti throat and breast wedge, 
six have complete midventral bisection of the red by an agouti line, 
and two have no, or virtually no, red. (5) There is a strong tendency 
for the feet to be either black or blackish agouti. 

Numbers 1, 2, and 5 above are characters widespread in the spe- 
cies flavimanus and emphasize this relationship. Numbers 3 and 4 
particularly emphasize relationship to subspecies castaneoventris, not 
to the geographically nearer ningpoensis. 

Callosciurus ferrugineus (F. Cuvier) 

Sciurics ferrugineus F. Cuvier, 1829 Table General et Methodique (appended 
to Geoffrey St. Hilaire and F. Cuvier, Histoire Naturelle de les Mammi- 
feres, 3, p. 238). 

Sciurus keraudreni Lesson, 1830, Centurie Zoologique, pi. 1 and text. 

Types. — S. ferrugineus, not found at MNHN in 1951, collected 
by Duvaucel "from India"; keraudreni, not found at MNHN in 1951, 
taken in Pegu, Burma. 

Material examined, all from Burma. — Mt. Popa, 4960 feet 
(AMNH), seven, (BM), four; "Kodugwe," Pegu Yoma (AMNH), 
three; "Yetho River," Pegu Yoma (AMNH), two; "Owegoo Dis- 
trict," Pegu Yoma (BM), one; Chaunglon, Tkyaulla District (BM), 
one; Daingmhu, 200 feet, 40 miles north of Pegu (BM), two; Gok- 
teik, 2133 feet, northern Shan states (BM), one; Kaing River, S. Py- 
inmana (BM), two; Lawksawk State (BM), one; "East India" 
(MCZ), two; 700 feet, northern Zamaya Reservation, 70 miles north 
of Pegu (BM), one; 300 feet, southern Zamaya Reservation, 60 miles 
north of Pegu (BM), one; Rangoon (MNHN), one, (BM), five; 
"Tankton," Rangoon (BM), one; 

Definition. — Callosciurus ferrugineus is here regarded as mono- 
typic (but including a named form keraudreni which may prove to 
occur west of the lower Irrawaddy and to be a good subspecies) and 
occurs only in Burma and principally between the Sittang Valley 
and the lower Irrawaddy. See the mapped distribution in Figure 14. 

Diagnosis. — The dorsal pelage of Callosciurus ferrugineus is all a 
glossy, rich, dark red except that it darkens to blackish on the feet, 
and that in the tip of the tail there is a small tuft of white hairs. The 
ventral pelage is entirely a lighter red than the back. These char- 
acters distinguish ferrugineus from all other species of Callosciurus. 


Relationships to other species. — It is obvious that the wholly red 
squirrels, ferrugineus, which live in the forests of the Pegu Yoma of 
Burma above Rangoon (see fig. 14) very closely resemble the wholly 
red squirrels of Thailand, Cambodia, and Laos. Nor can it be re- 
garded as surprising that earlier authors (Ellerman, 1940; Ellerman 
and Morrison-Scott, 1951) have placed them in a single species. 
Their present ranges as known to us from collected specimens, are 
some 200 miles apart (see fig. 14) and separated by one great river 
barrier, the Sal ween. (Geologically earlier these separately evolving 
red squirrel populations must have been separated also by the com- 
bined Irrawaddy and Sittang rivers, as will be discussed in detail 
below) . 

The orange-footed form C. erythraeus sladeni of upper Burma 
closely resembles the orange-footed C. /. flavimanus of Laos and Viet- 
nam and is separated from it by more miles and one more barrier 
river, the Mekong, than C. ferrugineus is from the all red [C. finlay- 
soni] menamicus of northern Thailand. But since C. e. sladeni and 
C /. flavimanus are shown above to be separately evolved, one might 
anticipate something of the same sort of separate origin of the all 
red ferrugineus from that of the all red menamicus, williamsoni, and 

Chhibber (1934) reports that the Chindwin and Irrawaddy were 
formerly quite separate rivers and that a small lateral branch of the 
Chindwin must have by head erosion performed an act of stream 
piracy which diverted the whole upper Irrawaddy north of Manda- 
lay westward into the Chindwin. The former lower course of the 
Irrawaddy River was clearly down the Sittang Valley almost straight 
south from Mandalay to the sea. See, for example, de Terra's (1944, 
p. 71) sketch map of the physiography of upper Burma. Chhibber 
(1934, p. 20) implies that this piracy occurred after certain geological 
events which "characterised the close of the Tertiary ..." 

Prior to the piracy performed by the Chindwin, then, the Chind- 
win and Irrawaddy held between them a narrow strip of land con- 
sistently about 100 miles wide and at least 650 miles long, from the 
Tibetan Plateau to the sea. In the upper 50 miles of this strip the 
squirrel species Callosciurus erythraeus is a gray agouti dorsally and 
red only on the venter (intermedius) . Southward from there the spe- 
cies descends from the mountains to forests of the wide valleys, and 
the pelage of the rostrum, feet, and tail are red (sladeni). The south- 
ernmost known specimens of sladeni, from Yin (about 22° 43' N.), 
are very red. 


It seems a most reasonable hypothesis that in the 400-mile length 
of this inter-riparian strip remaining south of Yin, the population of 
these squirrels became redder until entirely red. When the stream 
piracy was accomplished by the Chindwin, then, the wholly red pop- 
ulation would have become cut off completely from contact with any 
population with which it could interbreed. It seems probable that 
enough further differentiation has taken place in ferrugineus during 
this isolation that interbreeding would no longer take place if natural 
access to the adjacent population of sladeni were restored. No en- 
tirely red specimens are known from north of the new river barrier. 

While the stream piracy barred ferrugineus from spreading to the 
north, it opened an area to the east across the empty former bed of 
the Irrawaddy. It is interesting to note that ferrugineus has spread 
across this barrier very little. Of the 13 collecting localities from 
which we have examined specimens, only two are across this former 
barrier: Gokteik and Lawksawk. No specimens which might be in- 
tergrades between ferrugineus and either shanicus or phayrei are 
known to us, although specimens of all three forms are now known 
from the single collecting locality of Gokteik. No evidence at all is 
known that C. flavimanus shanicus has crossed the former barrier to 
the west. Possibly its ecology is too specialized. Better details from 
the field on both geographical and ecological distribution of ferru- 
gineus, phayrei, shanicus, and janetta on both sides of the broken 
barrier would be interesting. 

Pelage color. — The dorsal pelage is generally a glossy Morocco 
Red, mid-dorsally deepening to about Claret Brown. The tail is 
Maroon, and within its tip a small cluster of white hairs regularly 
occurs. Ventral pelage is also red but a little lighter, about Mahog- 
any Red in most specimens but as light as Burnt Sienna in some. 
The dorsal pelage color extends onto the feet but darkens near the 
digits which are almost black. It is the blackish feet and white tuft 
of hair in the tip of the tail that distinguish ferrugineus from the all 
red squirrels of the species finlaysoni. 

Discussion. — Lesson's plate of keraudreni indicates a much greater 
length of the white tail tip than is the case in ferrugineus. There is 
in the British Museum a specimen. No., which is labeled 
"keraudreni." It is from Arakan in west Burma, and this may pro- 
vide a notable westward extension of the species. Almost two inches 
of the tip of its tail are white. It thus agrees substantially with Les- 
son's plate and differs somewhat from true ferrugineus in which the 
white tip is very much smaller. Confirmation of the occurrence of 


dark-footed, all red, squirrels occupying a geographic area east of 
the present lower course of the Irrawaddy in forest of the Arakan 
district, especially if the long, white tail tip is characteristic of it 
there, would justify recognition of keraudreni as a subspecies of fer- 

Nothing that the writer has seen in southern material of C. eryth- 
raeus erythrogaster suggests intergradation farther south with a red 
subspecies. However, no barrier is known to prevent such inter- 
gradation now or formerly, and the area in the Arakan Yoma from 
which no specimens have been studied and whence evidence of inter- 
gradation might yet be obtained, is quite large (fig. 13). If there is 
a geographic population of red squirrels south of the range of erythro- 
gaster, its true relationships will remain uncertain until someone can 
show evidence that it does or does not intergrade with erythrogaster. 
Lacking such evidence, we are assuming that if a red population does 
live there, it became transferred west of the main barrier of the Irra- 
waddy in the delta by a shift of the main flow to a more eastern chan- 
nel, and subsequent silting up of the old channel until it could be 
crossed through vegetation by squirrels. We recognize that species 
level differentiation of C ferruginens from C. erythraeus is tenuous 
and hypothetical. 

Field observations by an experienced field collector are available: 
"On Mt. Popa this species only occurred in the thick jungle on the 
higher slopes of the mountain. Seen from a distance in a tree it ap- 
pears black, the white tail tip showing up conspicuously." (G. C. 
Shortridge in Wroughton, 1915a, p. 473.) 

Callosciurus finlaysoni (Horsfield) 

Definition. — The species Callosciurus finlaysoni is constituted by 
subspecies finlaysoni, folletti, trotteri, frandsoni, alhivexilli, harmandi, 
germaini, nox, cinnamomeus, annellatus, williamsoni, menamicus, sin- 
istralis, bocourti, and boonsongi (and see their synonymies). This 
species occupies the forests of Thailand, Cambodia, and the part of 
Laos along the Mekong as shown in Figure 14. 

Diagnosis. — There are populations of this squirrel with entirely 
white pelage, other populations with entirely black pelage, and still 
others with entirely red pelage, and there is no single character now 
known to be diagnostic for the species. Table 8 shows the dimen- 
sions of some type specimens of forms here included in the species 
finlaysoni, and the color characters are discussed in the subspecies 

1 ^ Q3 / Si { 1 

+ 22-H 

+ 20" N 


ir^fl _r.©<^ 



Fig. 14. Geographic distribution of two species, the Pegu red squirrel, Cal- 
losciurus ferrugineus, A, and the Thailand tree squirrel, Callosciurus finlaysoni, 
B to P, as plotted from material examined. Subspecies of finlaysoni: B, finlaysoni; 
Cfolletti; D, trotter; E, frandseni; F, albivexilli; G, hardmandi; (off the S. E. corner 
of the map about 80 miles SSE on a small island is) germaini; I, nox; J, cinna- 
momeus; K, annellatus; L, williamsoni; M, menamicus: N, sinistralis; 0, bocourti; 
and P, boonsongi. 



Relationships to other species. — There is in Thailand, Cambodia, 
and Laos a complex of squirrels so extraordinarily differentiated that 
their variation has to be seen to be believed. Judging by collections, 
various local populations of it exhibit pelage that is entirely white, 
other populations entirely red, others entirely black, black above 
with white below, and other more complicated patterns. About 28 
to 30 subspecies names have been applied to color variations of this 
presumed species. 

After having studied the outstanding collection of squirrels of this 
complex in the United States National Museum listed in the follow- 
ing pages, and having plotted as carefully as we could the many col- 
lecting localities of it, we feel that this has substantially advanced 
understanding of these squirrels, but that a great deal of mystery 
remains to be dispelled. We feel that we have improved knowledge 
of this complex not by discovering what its true relationships are, 
but by showing better how some of the problems need to be attacked 
locally in greater detail. 

Thus, even though we accept finlaysoni here as a species, it is pro- 
visionally and with some serious reservations. Its red subspecies 
williamsoni may quite well intergrade with C flavimanus flavimanus, 
but evidence that it does is scanty (see figs. 13 and 14). There are 
indications that C. finlaysoni sinistralis crosses (or intergrades?) with 
C. flavimanus thai, but just north of the range of thai, sinistralis 
seems to occur sympatrically with C. flavimanus zimmeensis or to in- 
terdigitate with it in a substantial area where no evidence of inter- 
breeding has come to our attention. 

Callosciurus finlaysoni is one of the species reported earlier to 
possess but two pairs of functional mammae (Moore, 1961a, p. 14) 
before the present revision of this species was accomplished, and it 
may be worth mention that no exceptions to this were observed in 
the larger series at the United States National Museum during this 
revision. The following two records of brood size hint that in finlay- 
soni brood size may also be small. Bonhote (1901b, pp. 53, 54) notes 
one specimen from above Uttaradit on the Mae Nan that would now 
be known as subspecies menamicus, which was taken "pregnant with 
two young" on April 3, 1900; and another specimen which would 
now be known as subspecies sinistralis which was "pregnant with 
two young" when taken on February 3, 1900, at Kamphaeng Phet 
on the Mae Ping. 

This species finlaysoni is for the most part a tree squirrel of the 
lowland forests. It is rather small, and it shares nearly all of its 








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range with another species of generally lowland tree squirrel, Callo- 
sciurus caniceps which is a somewhat larger animal. The latter does 
not appear, however, to be sympatric with the finlaysoni subspecies 
cinnamomeus, annellatus, and ivilliamsoni. 

Callosciurus finlaysoni finlaysoni (Horsfield) 

Sciurus finlaysoni Horsfield, 1824, Zoological Researches in Java, No. 7, sev- 
enth unnumbered page of text. 
Sciurus finlaysoni portus Kloss, 1915, Jour. Nat. Hist., Soc. Siam, 1, p. 158. 

Types.— S.f. finlaysoni, BM No., old male from Koh 
Si Chang, Gulf of Siam, collected by Greorge Finlayson; portus, BM 
No., adult male from Koh Si Chang collected January 26, 
1915, by C. B. Kloss. 

Material examined. — Koh Si Chang, Gulf of Siam (CNHM), two; 
(BM), nine, (USNM), 22. 

Pelage color. — Description from Kloss (1915a, p. 158) : "Ivory yel- 
low throughout, most intense on the upper surface and tail but paler, 
almost white, on sides of head, chin, and forelimbs. Skin of nostrils, 
lips, ears, soles of feet, and genitals black. Eyes black. Vibrissae 
black" (in the type but see below). 

Discussion. — Wroughton (1908, p. 398), as first reviser of finlay- 
soni, stated that "The type locality is the island of Sichang." Kloss 
(1915b) ignored this as if a matter of opinion and tried to show that 
specimens of finlaysoni originally obtained on the mainland of Siam 
were intended by Horsfield to represent the new species and that 
mention that one specimen of the type series was from Koh Sichang 
was purely incidental. In consequence, he named the Koh Si Chang 
squirrel portus. Robinson (1916, p. 35) points out that Wroughton's 
restriction of the type locality to the island of Sichang is type desig- 
nation by the first reviser. Robinson has been followed by other 
authors on this matter, including Kloss (1919, p. 368). 

Habits. — Kloss's (1915a, p. 158) remarks seem well worth quot- 
ing: "I found S. f. portus very common on Koh Si Chang and very 
fearless. Its small size is due to its insular habitat and to the poverty 
of the vegetation, large scrub rather than forest, with which the island 
is covered. It runs busily about the stems and branches, often al- 
most descending to the ground, in its search for food. The pelage 
of many of the animals obtained is considerably abraded and in every 
case has a sticky feeling very different from the smooth glossy hair of 
other squirrels, while both as skins and in the flesh a peculiar char- 
acteristic odour attaches to them. 


"In the series a number of animals have black vibrissae like the 
type, in an equal number the vibrissae are pure white, while some 
have them black and white mixed. In about half the series the hairs 
of the distal half of the tail are black tipped." 

Callosciurus finlaysoni folletti (Kloss) 

Sciurus finlaysoni folletti Kloss, 1915, Jour. Nat. Hist. Soc. Siam, 1, p. 159. 

Type.— BM No., old female from Koh Phai, an island 
15 miles south of Koh Si Chang, Gulf of Siam, taken February, 1915 
by collectors of C. B. Kloss. 

Material examined. — Koh Phai, Gulf of Siam (USNM), 12, 
(CNHM), 1, (BM), 12. 

Pelage color. — A small white squirrel, the hairs white-tipped 
throughout, but with bases gray, the gray showing through the 
white to give an appearance of dirty white or even grayish white. 
This can at best be only very slightly different from the form on Koh 
Si Chang. A tendency is seen for a wash of red to appear on under 
parts, especially in females. 

Callosciurus finlaysoni trotteri (Kloss) 

Sciurus finlaysoni trotteri Kloss, 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 178; 
ibid, 3, p. 369. 

T?/2?e.— USNM No. 236601, adult male from Koh Lan, an island 
10 miles south of Koh Phai, Gulf of Siam, collected by C. B. Kloss 
on October 29, 1916. 

Material examined. — Koh Lan, Gulf of Siam (USNM), 9; 
(CNHM), 1. 

Pelage color. — This is a white-tailed, gray-bodied, squirrel with 
blackish extremities and a rather faintly expressed atrodorsalis mark 
(the posterior two- thirds of the mid-dorsum blackish in a band 30 
or 40 mm. wide). The gray dorsal pelage is composed of hairs that 
are whitish (Pale OHve Buff) on their distal half and neutral gray 
basally. The basal gray shows through. The limbs and sides of 
head and neck are gray slightly tinged with brownish, and the limbs 
darken distally to slightly agouti blackish hands and feet. The ven- 
tral pelage is also grizzled gray but has whitish inguinal and axillary 
spots. The tail is whitish above with an inner tuft of hairs among 
the especially long hairs of the tail tip black, and whitish below ex- 
cept for the very bases of the hairs which are blackened in most of 
the specimens. 


Callosciurus finlaysoni frandseni (Kloss) 

Sciurus ferrugineus frandseni Kloss, 1916, Proc. Zool. Soc. London, 1916, p. 46. 

Type.— BM No., adult male from the island Koh 
Chang, southeast Thailand, collected December 12, 1914, by C. B. 

Material examined. — Koh Chang, southeastern coast of Thailand 
(USNM), nine, (BM), four. 

Pelage color. — This is a red squirrel with gray sides. It is red on 
top from rostrum to tip of tail, and red throughout the ventral pelage, 
but the sides of the body, head, neck, and dorsal pelage of the legs 
are an agouti of brownish olive. The feet are red. The anterior 
part of the throat is gray. 

Some of the mainland material included under cinnamomeus is 
quite like frandseni, and it may be that further collecting will show 
that the Koh Chang material and that on the mainland like it should 
be recognized as a single subspecies different from cinnamomeus. 
See further discussion of this under the subspecies cinnamomeus. 

Callosciurus finlaysoni albivexilli (Kloss) 

Sciurus albivexilli Kloss, 1916, Proc. Zool. Soc. London, 1916, p. 47. 

Type.— BM No., old male from Koh Kut, an island 
off the southeast coast of Thailand, collected December 25, 1914, 
by C. B. Kloss. 

Material examined.— Koh Kut, Thailand (BM), 7, (CNHM), 2, 
(USNM), 11. 

Original description. — ". . . Black throughout except the extrem- 
ity of the tail, which is white . . . [This subspecies] is somewhat vari- 
able in respect of the white tail tip. In some animals the last 3 or 
4 inches of the tail are white ... in others . . . the pale tip is reduced 
to a bunch of grayish hairs at the extreme end ..." 

Discussion. — Only the constantly present (though variable in ex- 
tent) white tip of the tail distinguishes this insular subspecies from 
nox. No taxonomic difference is seen between the skulls of the two 

Callosciurus finlaysoni harmandi (Milne-Edwards) 

Sciurus harmandi Milne-Edwards, 1876, Bull. Soc. Philomathique, Paris (ser. 6), 

13, p. 8. 
Callosciurus ferrugineus pierrei Robinson and Kloss, 1922, Ann. Mag. Nat. 

Hist. (ser. 9), 9, p. 91. 


Types.S. harmandi, MNHN No. 1876-125 (140), from Phu 
Quoc, south of Cambodia, collected by Harmand in 1876; pierrei, 
BM No., an old female taken on Phu Quoc, an island off 
the coast of Cambodia in February 1874 by Pierre. 

Material examined. — Phu Quoc, Cambodia (MNHN), 3, (BM) 

There are three paratypes of harmandi. The skulls, from which 
many of the teeth have fallen, of the four animals were not marked 
by number. One of us (Tate) tentatively marked numbers on all 
four (including the type) from one to four. There seems to be no 
possibility that they can ever be matched to their proper skins. 

Type description. — The color of the pelage of the type (examined 
in 1951) is dark reddish brown, quite heavily grizzled with whitish 
posterior to the nape, but the top of the head is dark brown. The 
tail appears whitish in over-all views, but the bases of the hairs are 
very dark gray, and the tail tip is dirty white. The under parts are 
light orange red, with bases of the hairs gray, but on the chest the red 
gives place to buff, and the throat is dark gray brown like the sides 
of the head. The relationships of harmandi appear to be with sini- 
stralis, although harmandi is geographically remote from it and a dis- 
tinctly insular race. 

Discussion. — The alleged presence of two kinds (harmandi and 
pierrei) of red-bellied squirrels on Phu Quoc poses something of a 
problem. In the first place, the known materials representing the 
two named forms, although very scarce, are admittedly different 
enough from forms on the adjacent mainland and from each other 
to be good subspecies. However, both of them seem definitely re- 
lated to the species finlaysoni in their color characters, and two sub- 
species of a single species do not retain characters distinct from one 
another while confined together on an island only 30 miles long. 

There seem to be at least two reasonable explanations: (1) The 
material described as harmandi and pierrei really all represents but 
one subspecies which is exceedingly variable. It is well documented 
that squirrels may be locally this variable (Moore, 1956). (2) The 
island has been invaded twice, and during the time interval between 
invasions the first emigrants and the nearest mainland form evolved 
so differently in appearance and habits that when contact was re- 
newed during the second invasion no interbreeding took place and 
competition has somehow been minimized. In this case it would 
probably be better to regard harmandi as a distinct species and pierrei 
as a subspecies of finlaysoni. 



New collecting or, at the very least, documented field study is 
needed on Phu Quoc to determine which of the two above conditions, 
or whether some other, prevails. Since double invasion seems to us 
the less likely situation, we have tentatively treated harmandi and 
pierrei as if they represent a single variable subspecies. 

Gallosciurus finlaysoni germaini (Milne-Edwards) 

Sciurus germaini Milne-Edwards, 1867, Rev. Zool., Paris (ser. 2), 19, p. 193. 

T^pe.— MNHN No. 1865-429 (126), old adult from Pulo Condor, 
Cochin China, collected by R. Germain. 

Material examined. — Pulo Condor, Cochin China (BM), 2, 
(MNHN paratypes), 10; "Cambodia" (BM), 1. 

Type description. — Though the color of the typical series appears 
at first jet black, a trace of brown mixed with the black of the chin, 
neck and under parts could be distinguished in good light when ex- 
amined in 1951. Possibly this is due to fading, as the specimens 
were collected nearly 90 years ago. In the type specimen there is 
also a tiny patch of hairs with whitish tips asymmetrically placed on 
the left of the lumbar region. Hands, feet and tail are perfectly black. 

Dimensions. — This insular subspecies apparently may be smaller 
than the black mainland subspecies about Sriracha, Thailand, from 
Wroughton's (1908, pp. 397-398) comparative data on an adult of 
each: hind foot lengths respectively 42 compared to 51 mm.; greatest 
skull lengths respectively 47 to 53.7; greatest zygomatic breadth, 
28 to 32 mm. 

Gallosciurus finlaysoni nox (Wroughton) 

Sciurus nox Wroughton, 1908, Ann. Mag. Nat. Hist. (ser. 8), 2, p. 397. 

Type. — BM No., adult male from 50 miles southeast of 
Bangkok, coastal Thailand at sea level, collected August 7, 1906, 
by T. H. Lyle. 

Material examined, all from the mainland coast of Thailand south- 
east of Bangkok. — Sriracha, at sea level (BM), three, (NR), one, 
(USNM), four; Hup Bon [Ban Hup Bon], 500 feet (BM), two, 
(USNM), one; Satahip (BM), two, (CNHM), two, (USNM), six; 
60 miles southeast of Bangkok (BM), two; Nongkhor [Ban Nong 
Kho] (USNM), 11, (MCZ), two; Ban Sadet (USNM), three; Nong 
Yang (USNM), one; Huey Yang (USNM), one. 

Discussion. — It seems possible that nox represents a color phase 
of cinnamomeus which has evolved to a high proportion of the pop- 


ulation through natural selection, possibly in dark mangrove forest. 
Specimens of cinnamomeus have been taken at five of the eight above 
localities, but at Sriracha and Satahip we have records of only black 
squirrels. It is worth noting that some selection for blackness may 
be taking place in the "red phase" in this area, too, since black on 
the feet and tail is the basis for the named form herherti from this 

Pelage color. — Description of nox is simple; it is jet black above 
and below. 

Callosciurus finlaysoni cinnamomeus (Temminck) 

Sciurus cinnamomeus Temminck, 1853, Esquisses Zoologiques sur la Cote de 

Guine, p. 250. 
Sciurus splendens Gray, 1861, Proc. Zool. Soc. London, 1861, p. 137. 
Callosciurus ferrugineus herherti Robinson and Kloss, 1922, Ann. Mag. Nat. 

Hist. (ser. 9), 9, p. 90. 

Types. — S. cinnamomeus, RNH No. 13367 (Temminck's specimen 
"C") adult from Cambodia; splendens, BM No. (= Gray's 
variety 2), adult female from Cambodia, taken by Mouhote; her- 
herti, CBK No. 2017 (not found at BM), an adult male collected at 
Hup Bon, near Sriracha, southeast Thailand, on July 25, 1915, by 
a museum collector. 

Material examined, from Thailand. — Ok Yam (USNM), four, 
(BM), four; Klong Yai (USNM), eight, (BM), two; Klong Menao 
(USNM), one, (BM), one; Lem Ngop (USNM), two, (BM), two; 
Ban Sadet, Sriracha (USNM), one; Nongkhor, near Sriracha 
(USNM), two, (CNHM), one, (MCZ), one; Lem Sing Mountain, 
Chantaboon (USNM), eight; Chantaboon (USNM), four, (BM), 
one, (UMMZ), two, (AMNH), seven, (ANSP), four; Kao Seming, 
Krat (USNM), six; "Bantam Dam" (USNM), one; Kao Sabab 
(USNM), 12, (CNHM), two; Nong Yang (USNM), two; Hoopbon 
[Ban Hup Bon] (USNM), two; "Huey Yang," Sriracha (USNM), 
seven; Sakeo, near Krabin (USNM), five; Khao Soi Dao, 3500 feet, 
S.E. Siam (ANSP), three; Kratt [Ban Krat] (ANSP), one; "Cochin 
China" (MNHN), two. 

Material examined, from Cambodia. — Bokor, 3000 feet (BM), 
five, (MNHN), three; Kampot, 100 feet (BM), one; Sien-reap, 150 
feet (BM), four, (MNHN), two. 

Pelage color. — In the United States National Museum collection 
of 47 specimens of this subspecies from southeastern Thailand, four 
general localities that are represented lie on a northwest-southeast 



axis. The southeastern group of localities (Ok Yam, Klong Yai, and 
Klong Manao) is at the Cambodian border, and the northwestern 
group (Hoopbon, Ban Sadet, Nong Khor, Sriracha, and Nong Yang) 
is near the east coast of the Inner Gulf of Siam about 50 miles from 
the mouth of the Chao Phraya, These two groups of collecting local- 
ities are represented by samples of 13 and 12 specimens respectively. 
The more southern of the two middle groups (Lem Ngop and Kao 
Seming) is represented by seven specimens, and the other (Chanta- 
boon, Kao Sabab, and Lem Sing Mt.) by 24. In the sample from the 
most southeastern group of localities 12 of the 13 possess all red pel- 
age. The other specimen is red except for olive agouti dorsal pelage 
of the cheeks, forelegs, and thighs. Of the next most southeastern 
seven only three are in the all red category; the other four are red 
except for olive agouti pelage on cheeks, forelegs, and thighs. The 
next northwestern sample, of 24, has 11 all red; 10 all red except for 
agouti cheeks, forelegs, and thighs; and three that are all red except 
for agouti side of head, neck, and body, and agouti fore and hind legs. 
The most northwestern of the four groups of localities is represented 
by nine all red specimens; two with all red pelage except for agouti 
cheeks, forelegs, and thighs; and one with all red pelage but agouti 
side of head, neck, and body, and agouti fore and hind legs. 

Discussion. — Consideration of the above information suggests the 
general distribution of an all red form of squirrel throughout this 
southeastern area of Thailand with a general tendency to produce 
variants marked by one of two similar patterns of agouti pelage. The 
greatest incidence of agouti pelage in these mainland localities occurs 
in the two middle groups which are close to the island, Koh Chang. 
From that island the United States National Museum sample of nine 
is composed of two with all red pelage except for agouti cheeks, fore- 
legs, and thighs, and seven all red but for agouti sides of head, neck, 
and body, and agouti fore and hind legs. No entirely red specimens 
are known from Koh Chang, and the sample from there is entirely 
like the two agouti variants on the mainland and preponderantly like 
the one with the greater amount of agouti. The insular population 
is here admitted as a separate subspecies, frandseni, and the mate- 
rial from the two middle groups of mainland localities is considered 
intermediate between cinnamomeus and frandseni. Koh Chang is 
very close to the mainland and separated from it by quite shallow 

The important sample of five of these squirrels from "Sakeo, near 
Krabin" is not included in the above discussion, and their locality is 


well north of those discussed above, (It is shown above the "J" in 
Figure 14.) In a general way each of these squirrels has all red pelage 
excepting for agouti sides of head, neck, and body, and agouti fore 
and hind legs. Furthermore, one specimen has an almost completely 
olive gray agouti back and the tail hairs banded with black near the 
base of the tail. Thus, the incidence of agouti in this sample is a 
hundred per cent, and the agouti is also more extensive per individ- 
ual even than in the sample of frandseni. 

The importance seen in the "Sakeo, near Krabin" series, how- 
ever, is not in the inexplicably frequent and extensive occurrence of 
agouti pelage, but other characters. No. 253428 has rather whit- 
ish agouti sides resulting from very elongate subterminal white bands 
on the hairs. No. 253425 has very whitish ventral pelage. This is 
whitish agouti on chin and throat, very pale buff on the inferior sides 
of the legs, and buffy on the venter. There are also occasional all 
white hairs on the tail. These whitish aberrations in two specimens 
of five are, in view of the proximity of bocourti, considered to consti- 
tute evidence of interbreeding between cinnamomeus and bocourti. 

In regard to the wholly red specimens of cinnamomeus, they are 
generally glossy and possess a broad, longitudinal, middorsal band 
that is notably darker than the red of the sides and legs. This is 
characteristic of the type of splendens, and distinguishes most cin- 
namomeus from menamicus (with which cinnamomeus is not in geo- 
graphic contact). There are occasional specimens of cinnamomeus, 
however, that are all red and that have middorsal pelage the same 
color as that of the sides and legs, and with less gloss to the pelage. 
These are the characteristics of the type specimen of cinnamomeus. 
Examples are USNM No. 255743 from Lem Sing, 257734 from Huey 
Yang, and 256840 from Nong Yang. 

We have seen in the small amount of material from Cambodia 
no evidence of intergradation between cinnamomeus and annellatus. 
But the white ring about the basal end of the tail in annellatus is the 
principal difference between it and all red cinnamomeus. Another, 
slighter difference is that in a specimen of annellatus the pelage of 
the back is the same red as that of the sides and venter. It would be 
surprising, indeed, if these two forms did not intergrade. 

Callosciurus finlaysoni annellatus (Thomas) 

Callosciurus ferrugineus annellatus Thomas, 1929 (for 1928), Proc. Zool. Soc. 
London, 1929, p. 839. 


Type.— BM No., adult female from Ankor, 150 feet, 
Cambodia, collected December 24, 1927, by J. Delacour and W. Lowe. 

Material examined. —Angkor, Siem Reap, Cambodia (USNM), 
one, (BM), one, (MNHN), one; Bassac, west of Mekong R., Laos 
(CNHM), two; "Cambodia" (CNHM), one; Sambor, 200 feet, Cam- 
bodia (BM), one; east of Paks^ at 1000 feet, Laos (BM), one. 

Pelage color. — The Chicago Natural History Museum specimens 
listed above are as follows: The dorsal pelage is Morocco Red, the 
ventral pelage is Claret Brown in two but Morocco Red in one, the 
tail is Victoria Lake with a sharply contrasting band of Cream Color 
about four centimeters long that begins about five cm. from the body. 
The face, feet, and wrists are darker than Maroon, 

Discussion. — In describing the subspecies annellatus, Thomas 
(1929, p. 840) notes that it is the "variety 1" ascribed by Gray (1861) 
to splendens, and which Thomas (1929) re-examined at the time 
(BM specimen no. Thomas (1929) regards Wroughton's 
(1908, p. 397) statement, "I have concluded to accept Gray's splen- 
dens var. 2 Capt. Flower's Chataboon specimen ... as the normal of 
the species." as a lectotype selection. The examination of further 
material now makes the choice seem a quite acceptable one, and we 
certainly concur in it. 

Callosciurus finlaysoni williatnsoni (Robinson and Kloss) 

Callosciurus ferrugineus wilUamsoni Robinson and Kloss, 1922, Ann. Mag. 
Nat. Hist. (ser. 9), 9, p. 90. 

Type.— BM No., adult female taken at Khet Don 
Heng, below Xien Khan [Chieng Khan] on the north bank of the 
Paklay Loop of the Mekong River on January 31, 1920, by H. C. 
Robinson and C. B. Kloss' collector. 

Material examined, all from Laos. — Vientiane (USNM), one; 
(MCZ), one; "Tha Ngon," Vientiane (CNHM), six; Paks^ (CNHM), 

Discussion. — The describers of this subspecies possessed a hy- 
podigm of 17 specimens from localities spread 200 miles along the 
Laotian bank of the Mekong River from Muong Liep (about 30 miles 
up river from Pak Lay) to Ban Manao (longitude 104° E.). Four 
specimens reported here from Paks^ extend the range southward 
along the east bank of the Mekong another 200 miles. It seems to 
be unknown from the other side of the Mekong. 

At Paks^ the ranges of wilUamsoni and annellatus seem to ap- 
proximate, and annellatus, curiously, is known from both sides of the 


lower Mekong. Perhaps this could have come about by naturally 
changing channels where the river is strongly braided beginning about 
50 miles south of Paks^. 

As is evident from material cited above, collecting in Laos has 
been meager, and from the little material available, scarcity of evi- 
dence on intergradation need surprise no one. There is a specimen 
of flavimanus (USNM No. 254753) from "Nong Kai, Laos," and if 
this locality may be taken to mean Laos, across the river from Nong 
Kai, Thailand, then this specimen is within the range of C. finlay- 
soni williamsoni. Possibly the Nong Kai specimen is, and certainly 
the one from the Bolovens Plateau (AMNH No. 87431) seems to be, 
a cross between C. flavimanus and C. finlaysoni williamsoni. 

Pelage color. — The following description is taken from Chicago 
Natural History Museum materials. The Tha Ngon series (taken 
June 30 to July 3) have dorsal pelage that is entirely Mars Orange 
except the tail tip which is faded. Ventral pelage is a darker color. 
Chestnut, than the dorsal pelage, except for new pelage in nos. 32385 
and 32390 which are molting, and 32388 which seems to have molted 
in parts. The Paks^ series (three taken November 18 and one 
taken January 2) have dorsal pelage entirely Morocco Red in three 
and Mahogany Red in the other (a November one). The tails are 
Maroon and very little faded at the tip. The ventral pelage is a 
lighter color, Mahogany Red, than the dorsal pelage in two and a 
still lighter color in the other two, Sanford's Brown. 

Discussion. — The above shows an early summer condition of 
darker ventral pelage than dorsal, and an early winter condition 
of the reverse. The molt mentioned above, which changes the ven- 
ter from Chestnut, which is darker than the back, to Orange Rufus, 
which is lighter than the back, suggests seasonal alternation of rela- 
tive intensities. The original describers' hypodigm was taken in 
January and February, and in it the venter is darker than the dor- 
sum. This suggests that a second seasonal molt (in early January?) 
may have returned the intensity relationship to the condition lost in 
the June-July molt. Probably no such neatly seasonal molting will 
be found to exist when more is learned. The molting of C. caniceps 
caniceps is not seasonal in the available museum material. 

The United States National Museum has a parous female speci- 
men, no. 240512, taken July 2nd at Luang Prabang, Laos, which 
Osgood (1932, p. 280) puzzled over but did not name. He assumed 
that it must belong to some species with which he was not working 
and not familiar, such as caniceps or pygerythrus. It is definitely not 


related to those, however, and examination of the evidence may show 
that it represents a cross between two forms with which he was then 
quite familiar. The specimen has an all red venter, an all red tail, 
and red middorsal pelage from snout to tail in a broad band covering 
the crown and pinnae but descending no lower on the sides. The 
remainder of the squirrel is cool gray agouti (sides, legs), but on the 
feet the gray agouti is rather strongly blackened. The red dorsum 
and tail are identical with williamsoni of Vientiane. The gray sides, 
the red of the venter, and the distinctively blackish agouti feet are 
like those of C. flavimanus hendeei of this vicinity. (C. /. hendeei is 
shown to intergrade with C /. flavimanus perhaps 40 air miles east 
of Luang Prabang in the above account of the latter.) The other 
species of Callosciurus east of the Mekong in this vicinity is C. inor- 
natus. See the ranges of hendeei and inornatus as "R" in Figure 13 
and triangles in Figure 15, respectively. The agouti of the sides and 
legs is more like that of hendeei than inornatus, the blackish feet wit- 
ness strongly against inornatus which has plain agouti feet. In un- 
damaged skulls of inornatus the ectopterigoid ridge skirts the outside 
of the foramen ovale like a little fence, but in hendeei and williamsoni 
and the specimen in question, the ridge runs directly toward the fora- 
men ovale and ends at its margin. 

Across the Mekong to the west among the normally all red C. fin- 
laysoni menamicus in Thailand are found specimens possessing the 
same pattern of red and agouti. See subspecies "M" in Figure 14. 
A series of these are USNM Nos. 261071 to 261074 and 267212 from 
Pua, Doi Phu Kha (2 specimens). Ban Na Ko, and Doi Kang Ma, 
respectively. These are further described below in the account of 
menamicus as crosses between C. finlaysoni menamicus and C. flavi- 
manus zimmeensis, and the Luang Prabang specimen differs from 
them in having the agouti pelage of the chin blend gradually with 
the red of the throat as in hendeei rather than extending onto the 
throat and breast as an agouti wedge partially bisecting the red 
venter as is common in zimmeensis. There is no indication of a sub- 
terminal black spot near the end of the tail in the Luang Prabang 
specimen, but the tip of the tail may be missing. The color of both 
the middorsal and the ventral pelage in the Luang Prabang speci- 
men is about Mars Orange, suggesting that the hendeei influence to 
lighter colored red venter altered the williamsoni pattern of ventral 
pelage differing notably from dorsal in intensity of color. 

In sum, the evidence indicates that the Luang Prabang specimen 
almost certainly represents a cross between C. finlaysoni williamsoni 


and C. flavimanus hendeei. This would indicate an extension of known 
range of williamsoni some eighty miles up the eastern bank of the 

Callosciurus finlaysoni menamicus (Thomas) 

Callosciurus ferrugineus menamicus Thomas, 1929 (1928), Proc. Zool. Soc. 
London, 1929, p. 839. 

Type.— BM No., an adult female taken from Nan, Thai- 
land, April 3, 1900, by Thomas H. Lyles. 

Material examined, from northern Thailand. — Den Chai (NR), 
one; Pak Koh (NR), two; Me Moh (BM), two; Uttaradit, 58 meters 
(BM), one; Lakhon, 235 meters (BM), one; Nan, 306 meters (BM), 
one; Ban Huai Som (USNM), two; Ban Me Mo (USNM), one; 
Pang Nam Un, Nan (USNM), two; Doi Pu Het (USNM), one; and 
Doi San Huai Wai (USNM), one. 

Material examined, of crosses or intergrades between menamicus 
and C. flavimanus zimmeensis from Thailand. — Pua (USNM), one; 
Doi Phu Kha (USNM), two; Ban Na Ko (USNM), one; and Doi 
Kang Ma (USNM), one. 

Diagnosis. — This subspecies may evidently be distinguished from 
other all red subspecies of the Indochinese Subregion by having the 
back, sides, venter, and feet entirely and equally red, and the tail 
red but with a buffy white tip. 

Discussion. — The intermediates formed between this subspecies 
and C. flavimanus zimmeensis have a subterminal black spot in their 
red tails and their red venters partly bisected anteriorly by a poste- 
riorly directed wedge of agouti pelage. All but the Doi Kang Ma 
specimen have red venters, backs, and tails, but agouti sides. 

Gyldenstolpe (1916b, p. 37) noted this red squirrel "to be fairly 
common in the dry forests of Northern Siam," and that "No white 
specimens were obtained or observed during the whole journey." 
He took only two specimens of this red form, both about Pak Koh 
where (Gyldenstolpe, 1916a, p. 5) he spent a month in March and 
April, and wrote that "The mixed dry forests are . . . the most pre- 
dominant in the low-lying country and on the lower hills." Deignan 
(1945, p. 25) agrees that the "Mixed-deciduous [forest is] an associa- 
tion highly developed at low elevations in the larger valleys of our 
[North Thailand] provinces . . ." 

The hypodigm of two specimens from Doi Souket on which the 
name primus (synonym of C. flavimanus zimmeensis) is based is, in 


view of the present knowledge of the geographic ranges of these 
squirrels, probably evidence of interbreeding between zimmeensis 
and menamicus. They are closer to the former, having only reddish 
ears and nape and red posterior half of the tail as suggestive evidence 
of relationship to menamicus. (The undivided venter may be found 
occasionally elsewhere in the sample of zimmeensis.) 

This interbreeding is between a tropical montane form {zimmeen- 
sis) which in this artenkreis is characteristically agouti gray-olive- 
brown dorsally, and a tropical lowland subspecies {menamicus) which 
in this artenkreis tends to be more brilliantly colored dorsally. It 
thus seems appropriate that the type of primus should be from a river 
bank at 1500 feet, but that the other specimen should be labeled 6500 
feet is surprising. (Deignan, 1945, p. 17, gives the elevation of Mt. 
Souket as only 5,958 feet, but it is not the disagreement of 500 feet 
that seems significant.) 

The United States National Museum specimen from Luang Pra- 
bang has a red dorsum from snout to tail, a red tail, and an undivided 
red venter, but the sides, legs, and feet are strictly agouti. The 
agouti sides are separated abruptly from the red dorsum and red 
venter alike. This specimen seems in color characters so much like 
the Pua and Doi Phu Kha intergrades between menamicus and zim- 
meensis that one could think it must have been taken on the west 
side of the Mekong opposite Luang Prabang. See the discussion of 
it above in the account of williamsoni. 

Callosciurus finlaysoni sinistralis (Wroughton) 

Sciurus bocourti sinistralis Wroughton, 1908, Ann. Mag. Nat. Hist. (ser. 8), 

2, p. 399. 
Sciurus bocourti dextralis Wroughton, 1908, Ann. Mag. Nat. Hist. (ser. 8), 2, 

p. 400. 
Sciurus bocourti lylei Wroughton, 1908, Ann. Mag. Nat. Hist. (ser. 8), 2, 

p. 401. 
Sciurus bocourti gruti Gyldenstolpe, 1917, Handl. Kungl. Svenska Vet. Akad., 

57, no. 2, p. 37. 

Types. — S. b. sinistralis, BM No., adult male from Kam- 
pong below Pichit (on Me Nam River), 35 meters, Siam, collected 
June 8, 1903, by T. H. Lyle; dextralis, BM No., young female 
from Kampeng [Phet] on the Me Ping, Thailand, taken February 3, 
1900, by T. H. Lyle; lylei, BM No., young adult female 
from Chiengmai, Me Ping River, Thailand, collected August 12, 
1907, by T. H. Lyle; gruti, NR No. 15, adult female from Bang Hua 


Pong on the southern slopes of Doi Khun Tan, north Thailand, col- 
lected on May 8, 1914, by Nils Gyldenstolpe. 

Material examined, all from Thailand. — Doi Lang Ka (USNM), 
one; Raheng, 110 meters (USNM), two, (BM), two; Kuhn Tan 
(USNM), eight, (ANSP), one, (NR), three; "B. Nam Kien," Nan 
(USNM), one; Pang Me Ton (USNM), two; Klong Klung (USNM), 
seven; Doi Phra Chao (USNM), six; Nakon Sawan (USNM), one; 
Wung Pratart Farm (CNHM), 16; 20 miles east of Chieng Mai 
(ANSP), one; 35 and 40 miles above Pichit (BM), two; Phitsanu- 
loke, 47 meters (BM), four; 160 miles north of Bangkok, 32 meters 
(BM), one; Bon Mee, Siang Hai, 90 meters (BM), one; "Banthee," 
Siang Hai, 90 meters (BM), one; above Kampeng on the Meeping 
River, 115 meters (BM), one; Meeping River, 65 meters, 180 miles 
north of Bankok (BM), one; Sokotai, 64 meters (BM), two; Bang 
Hue Pong (NR), three; Paknampo (USNM), one; Meping River, 
18° N. latitude, 310 meters (BM), one; south of Chiengmai, latitude 
18° 30' N., 260 meters (BM), one; Doi Mei Lai (USNM), one; Doi 
Me Kong Kha (USNM), two; Chieng Mai (USNM), two; Doi Nang 
Keo (USNM), one; Doi Chieng Dao (USNM), one; Doi Kang Ma 
(USNM), one; Pua (USNM), one; Doi Phu Kha (USNM), two; Ban 
Na Ko (USNM), one. 

Pelage Color. — The fine series of sixteen at Chicago Natural His- 
tory Museum from Wung Pratart Farm, Kamphaeng Phet Province 
have undivided Mahogany Red venters except for one being San- 
ford's Brown. On the head, wrists, ankles, and ears the pelage is red, 
about Hays Russet. The feet are an agouti of about Carob Brown 
(XIV). At first inspection the dorsal pelage does not appear to be 
agouti, but rather a random mixture of white hairs with red hairs and 
black hairs. Closer inspection reveals the "red" hairs to be black 
hairs with one or two reddish bands. The effect of the admixture of 
white hairs produces a more pronounced grizzle than one finds in 
most agouti pelages. The pelage of the back extends out the tail 
about 40 mm. Then there is a cream colored band 30 mm. wide 
across the tail, and beyond it the tail is pure Morocco Red in most. 
In some, however, the tail hairs are annulated black and white near 
their bases instead of entirely red. 

Diagnosis. — Near the base the tail is whitish for 30 or 40 mm, 
and beyond that Morocco Red, but the pelage of the dorsum varies 
from reddish agouti with many white or gray guard hairs to com- 
pletely gray pelage. 


Disctission. — The above pelage color description is very similar to 
Wroughton's original description of dextrcUis, but more detailed, and 
the locality is not far from the type locality of dextralis. 

Near the northern extremity of the range of the present subspecies 
at Khun Tan, ANSP No. 15301 has an undivided venter of Orange 
Rufous and dorsum of about Mouse Gray, and the tail, beyond the 
basal whitish ring (30 or 40 mm. long), is Morocco Red. The head 
and nape are Orange Rufous with a liberal sprinkling of gray guard 

The Khun Tan specimen described above seems possibly by its 
Orange Rufous head and nape, and geographical proximity to me- 
namicus, to represent intergradation with menamicus. USNM No. 
296943 from Ban Sanping, Nakhon Sawan Province, shows definite 
indication of crossing or intergradation with C. f. thai in having a 20 
to 30 mm. broad glossy black band on the posterior two- thirds of the 
middorsum, and otherwise possessing entirely sinistralis characters. 

There may eventually prove to be an adequate basis for breaking 
up this subspecies into two: sinistralis and lylei, the former distin- 
guished by the dorsal pelage containing black and /or brown, the 
latter by absence of brown in the grayish white dorsal pelage. How- 
ever, strong characters link the two potential forms and distinguish 
them jointly from surrounding subspecies. In absence of better 
knowledge, therefore, it seems desirable to recognize but a single 
subspecies here with three synonyms. 

Callosciurus finlaysoni bocourti (Milne-Edwards) 

Sciurus bocourti Milne-Edwards, 1867, Rev. Zool., Paris (ser. 2), 19, p. 193. 
Sciurus leucogaster Milne-Edwards, 1867, Rev. Zool., Paris (ser. 2), 19, p. 196 

(homonym of Cuvier, 1831). 
Sciurus leucocephalus Bonhote, 1901, Proc. Zool. Soc. London, 1901, p. 54. 
Sciurus floweri Bonhote, 1901, Ann. Mag. Nat. Hist. (ser. 7), 7, p. 455. 
Callosciurus finlaysoni tachardi Robinson, 1916, Jour. Federated Malay States 

Mus., 7, p. 36. 
Sciurus finlaysoni prachin, Kloss, 1920, Jour. Nat. Hist. Soc. Siam, 4, p. 103. 
Sciurus finlaysoni rajasima, Kloss, 1920, Jour. Nat. Hist. Soc. Siam, 4, p. 103, 
Callosciurus cockerelli Thomas, 1928, Ann. Mag. Nat. Hist. (ser. 10), 2, p. 100. 

Types.— Sciurus bocourti, MNHN No. 1860-123 (135), young adult 
from Thailand, collected in 1860 by Montigny; leucogaster, MNHN 
No. 1862-1241 (1303-A-224), adult male from Thailand; leucocepha- 
lus, BM No., adult male from Chainat, 20 meters. Me Nam 
River, Thailand, collected January 21, 1900; floweri, BM No. 99.2.- 


7.1, adult female from Klong Morn, near Bangkok, Thailand, col- 
lected August 13, 1898, by S. S. Flower; tachardi, BM No., 
old male from 75 meters, some 30 miles up the Mee Nan [river] above 
Uttaradit, 17^° N., Thailand, collected April 4, 1900, by T. H. 
Lyle; prachin, not seen. No. 2048 in private collection of C. B. 
Kloss, adult male from Krabin, central Thailand, collected Novem- 
ber 11, 1915, by E. G. Herbert and Malcolm Smith; rajasima, not 
seen. No. 2132 in private collection of C. B. Kloss, adult female from 
Lat Bua Kao, Thailand, collected October 10, 1916, by C. B. Kloss; 
cockerelli, BM No., adult from Paktoop Mountain, Nan, 
Thailand, collected January 1928, by Homer Wiesbecken. 

Material examined, all from Thailand. ^ — "Lam Ton Lang," Krabin 
(USNM), three; "Nong Mong" Muong Krabin (USNM), one; Kao 
Lem [Khao Laem] (USNM), one; Hin Lap (USNM), six; Pang Sok 
(USNM), one; Pak Chong (USNM), 28, (CNHM), two, (AMNH), 
15; Muek Lek (USNM), three; Nong Bua, Pasak River (USNM), 
one; Ban Manao Van, Pasak River (USNM), one; Manoram 
(USNM), one; Vichianburi (USNM), three; Bung Borapet (USNM), 
five; Nong Kai (USNM), one; Ban Den (USNM), one; Angton (BM), 
two; "Nong Dom" (UMMZ), seven; "Central Siam" (UMMZ), four; 
"Northeast Siam" (UMMZ), one; Lat Bua Kao (USNM), two; "Ban 
Den," on the Mekong (USNM), one; "B. Hin Ngom" (USNM), one; 
Prapoot Mtn., Lopburi (USNM), two; Ban Non Toulek, Chaiya- 
phum (USNM), one; Ban Lad, Chaiyaphum (USNM), two; "Ban 
Kudclon," Chaiyaphum (USNM), one; Pookeio, Chaiyaphum 
(USNM), two; Muaklek, Kaengkoi, Sraburi (USNM), one; "Tah- 
pen Mtn.," Muaklek (USNM), four; "Ban Na Nong," Chuempae 
[Ban Chum Phae] (USNM), three; "Sawan Mtn., Ban Seio," Loei 
(USNM), five; Ban Mung Ky [Muang Khai], Tahlee [Tha Li], Loey 
(USNM), 18; Lomlo Mtn., Ban Maeo Goksatawn, Dahnsai [Dan 
Sai], (USNM), 47; Lomloe Mtn., Ban Huie Muen, Goksatawn, 
Dahnsai [Dan Sai] (USNM), one; "Ban Na Muang, Na Haeo," 
Dan Sai, Loei (USNM), 23; Ban Muang Khai, Tha Li, Loei (USNM), 
16; Ban Nam Yen, Phak Khinak Mtn., Dan Sai, Loei (USNM), 25; 
Nam Lang Mtn., Ban Khok, Naphung, Dan Sai (USNM), 40; Ban 
Khana (USNM), one; Patoop Mountain, near Nan (BM), one; Nan 
(BM), one; "Tahkamen" (BM), two; Krabin (BM), one; latitude 
15° N., longitude 100° 30' E. (BM), one. 

The type locality, restricted to Ayutthaya by Wroughton (1908), 
is on the southern margin of the range of this subspecies, where its 
color characteristics are extremely variable. From the presently 


available information the range of hocourti extends from the vicin- 
ity of Ayutthaya northward up the Chao Phraya Valley and the Pa 
Sak Valley and beyond the latter valley's northern end to the banks 
of the Mekong. In the mountains on each side of the upper Pa Sak 
the large samples collected by Robert E. Elbel and deposited in the 
United States National Museum reveal constancy in those extreme 
characters which we find mark the geographic subspecies and which 
are but mildly expressed in the type specimen and hardly at all in 
some topotypes. 

Pelage color. — The 170 specimens of hocourti from eight localities 
near Dan Sai, Tah Li, and Loei about the headwaters of the Pa Sak 
are uniformly white or cream color on the entire venter, entire head, 
distal half of the dorsal pelage of the legs, lower half of the sides, 
ventral pelage of the legs and tail. The other color of this intensely 
pied squirrel is glossy black. The dorsum from just behind the ears 
and crown to a varying distance out the dorsal surface of the tail, 
and extending halfway down the sides and out half the length of the 
legs, is black. 

The black of the middorsum consists often of entirely glossy 
black hairs. The black pelage extending halfway down the sides 
generally is finely punctate with minute white bands on the black 
hairs. The finely punctate condition rather commonly occurs all 
over the back. Rarely (four instances in 170), the black dorsum 
possesses a liberal sprinkling of white hairs among the black, and a 
few white hairs occur infrequently among the others. The dorsal 
pelage of the tails is banded black and white, and in aggregate this 
gives the impression of 10 or 12 bars across the dorsal surface of 
the tail. 

The number of these 170 skins taken during each month of the 
year is: January, 16; February, 26; March, 39; April, 7; May, 19; 
June, 31; July, 0; August, 0; September, 3; October, 15; Novem- 
ber, 7; and December, 4. No seasonal variation in pelage color is 

Three nestlings were preserved as skins among this series of 170 
(February 18, March 24, and March 30) and from these, and the 
greater number of older immatures, it is evident that no regular 
ontogenetic change in the color pattern occurs. The black dorsal 
color of the nestlings is slightly less glossy and more brown than that 
of the adults. 

In specimens progressively more distant from this center of color 
pattern strength of the subspecies, the dorsum becomes variably 


and progressively paler until specimens which are entirely cream 
colored apparently predominate in peripheral populations (named 
ones of which are tachardi, prachin, and rajasima). 

Discussion. — It should surprise no one that so many variations 
of this variable subspecies have been named in the early exploratory 
phase of knowledge of these squirrels. Quite possibly there are one 
or more populations particularly of wholly cream colored animals 
which may yet merit recognition as geographic subspecies distinct 
from the pied one. In the present effort to consolidate knowledge 
in a revision and to offer a meaningful picture of the geographic 
variation, the possibly oversimplified interpretation of the available 
evidence given above seems best. 

In the south, specimens of bocourti have been collected near sta- 
tions along the railroad three fourths of the way from Ayutthaya 
toward Nakhon Ratchasima: the localities Hin Lap, Muak Lek, 
Pak Chong, and Lat Bua Kao. From these places bocourti is very 
nearly a wholly cream-colored form, but some are faintly gray dorsally 
in the pattern characterizing the intensely black-and-white bocourti 
from the north. The specimen from Kao Lem [Khao Laem] is one 
with a notably dark dorsum and from a locality a little south from 
Pak Chong. Farther south near Krabin Buri the ranges of bocourti 
and cinnamomeus must come together, and among the five specimens 
of cinnamomeus we have examined from Sakeo in that vicinity one 
provides clear indications of intergradation with bocourti in possess- 
ing ventral pelage that is nearly white instead of red, and a whitish 
eye ring like the type of floweri. 

On the west, the distribution of bocourti appears to be restricted 
by the Chao Phraya [river] across which it is opposed to C. flavimanus 
siamensis south of Ayutthaya. From Ayutthaya north to Nakhon 
Sawan C. flavimanus siamensis and thai and even C finlaysoni sinis- 
tralis intergrade or cross with one another across the Chao Phraya 
from bocourti. Whether crossing the river naturally or with the aid 
of man, bocourti seems to have injected some genetic material into 
the population of siamensis, for the exceedingly atypical white venter 
of the type specimen seems best explained by such a cross. North of 
Nakhon Sawan, bocourti seems from the records to occupy the east 
side of the valley along the large fork of the Chao Phraya called the 
Mae Yom, which separates the fairly white bocourti from the fairly 
dark sinistralis as far north as Muang Phitchit. Beyond Phitchit 
bocourti is replaced on the east side of the river by sinistralis (Wrough- 
ton, 1908, p. 399). No specimens evidently intermediate between 


these two forms have come to our attention. That intergradation 
between them may eventually be found seems suggested by the oc- 
currence of the sinistralis character, a 30-40 mm. ring of white on 
the tail near its base, observed to be clearly but not fully expressed 
in two United States National Museum specimens of intensely pied 
bocourti, one each from Ban Muang Khai and Ban Seio. 

On the north, both sinistralis and bocourti are replaced at Uttaradit 
and along the Mae Nam Nan by the wholly red form, menamicus. 
That all white bocourti do intergrade (or cross?) with the all red 
menamicus is to us clearly indicated by the intermediates (or hybrids) 
taken on Pahtoop Mountain above Nan and given the name cock- 
erelli. A specimen taken by Herbert Deignan from Ban Khana 
some 30 miles farther up the Nan Valley near Pua, is quite like the 
description of cockerelli in having pale gray sides of mixed white 
and agouti hairs, sharply marked of! venter, head and legs of cream 
color, but the broad stripe of Ferruginous from nape to base of 
tail along the mid-dorsum on the Pahtoop Mountain one is only 
pink in the Ban Khana specimen. Beyond the north end of the Nan 
Valley bocourti is probably replaced by C. flavimanus zimmeensis. 

The Mekong Plain of eastern Thailand has hardly been pene- 
trated by persons who collected mammals, and consequently the 
eastern extent of the range of bocourti for the most part remains 
uncertainly known. That it does extend out into the western edge 
of this plain is indicated by specimens taken at Ban Chum Phae 
by R. E. Elbel. That it does not occur all the way across to the 
eastern edge of the Mekong Plain is indicated by presence of another 
subspecies replacing it at Nakhon Ratchasima, and the few Thailand 
localities where tree squirrels have been collected farther east. 

We realize that this bocourti is a very strongly marked (even 
though peripherally such a variable) squirrel. We realize that it may 
in fact not intergrade with sinistralis, zimmeensis, or cinnamomeus, 
and that further collecting in the zones of bocourti contact with 
menamicus and these other forms may possibly reveal that only 
hybridization is taking place, and that bocourti is specifically dis- 
tinct from those forms. The evidence does not clearly indicate 
this now, however, and the more conservative course of retaining 
bocourti as a somewhat conglomerate subspecies seems the better 

To the east, bocourti probably does not progress to cream popu- 
lations, but undoubtedly intergrades with the subspecies named be- 
low, for in the collection from Nakhon Ratchasima one of the four 


(USNM No. 296516) is indistinguishable from bocourti and in the 
collection from Ban Sahng Kaw one of the seven (USNM No. 300061) 
is intermediate between bocourti and the following new subspecies, 
from eastern Thailand. 

Callosciurus finlaysoni boonsongi new subspecies 

Type.—VSNM No. 307801, an adult male taken on Phu Phan, a 
mountain in Sakon Nakhon District of Sakon Nakhon Province, 
eastern Thailand, June 14, 1954, by Robert E. Elbel and Boonsong 

Hypodigm. — Thirty-two specimens from Phu Phan (USNM Nos. 
307796 to 307827), the type locality; 12 specimens from Phu Do, a 
mountain in Nakae District, Nakon Phanom Province, eastern Thai- 
land (USNM Nos. 307828 to 307839) ; 10 specimens from Phu Kho, 
another mountain in Nakae District, Nakon Phanom Province, east- 
ern Thailand (USNM Nos. 307840 to 307849) ; seven specimens from 
Ban Sahng Kaw, Koek Pue District, Sakon Nakhon Province, east- 
ern Thailand (USNM Nos. 300061 to 300066 and 300072); four 
specimens from Nakon Ratchasima, formerly known as Korat, on 
the southwestern corner of the Mekong plain, Thailand (USNM Nos. 
296516 to 296519). 

Diagnosis. — This new subspecies differs consistently from bocourti 
in that the black or dark pelage of the back covers the sides down to 
the margin of the ventral pelage. (In bocourti with dark backs the 
cream color of the venter encroaches well up onto the sides.) 

Pelage color. — The color characteristics of this subspecies are com- 
plicated by occurrence in it of glossy black phase individuals (6 among 
the 65), creamy white phase individuals (3 among the 65), and red 
phase individuals (10 among the 65) . 

Ordinary color: These are almost wholly blackish squirrels, but vary 
much in the intensity of the black. Dorsal pelage is finely agouti 
and any color intermediate between black and a rather pale gray; 
always darker than the ventral pelage of the same individual. Ears 
regularly white-rimmed but occasionally all white, eyes sometimes 
ringed with white, but rostrum and feet rarely white or contrastingly 
lighter gray than the dorsum. Ventral pelage varying among indi- 
viduals but generally gray and most often longitudinally bisected by 
a line of darker pelage. The tail is generally blackish, but subtermi- 
nal white bands on the hairs occur in some individuals and become 
quite long posteriorly in still others. 


Reddish phase: Those in reddish phase include one male from Phu 
Kho, two adult males from Phu Do, four males and two females from 
Phu Phan, and one adult male from Ban Sahng Kaw. Most of these 
(7) have all red pelage on the ears, head, and venter, a little red on 
the tail distally, and considerable red infusion of the dark dorsal 
pelage. Others (3) have red ears, venter, and tail tip but are so dark 
a red as to be nearly black. 

Black phase: These possess entirely glossy black dorsal pelage except- 
ing whitish ears and inconspicuous subterminal white bands (about 
3 mm. long) on some hairs of the posterior half of the tail. The ven- 
tral pelage is blackish on the chin and throat, constituted by whitish- 
tipped, black hairs, and this extends posteriorly as a mid ventral 
wedge and then line bisecting the other ventral pelage. On each 
side of the blackish midventral line the ventral pelage is gray, and 
within it are axillary and inguinal paired spots of whitish pelage about 
12 to 15 mm. in diameter. (These include adults of both sexes and 
are one female from Phu Kho, two males and two females from Phu 
Phan, and one female from Ban Sahng Kaw.) 

Cream phase: This phase is exemplified by two females from Phu Do 
and one male from Phu Kho. They possess ventral pelage that is 
entirely cream color and dorsal pelage that has cream color pelage 
tips overlying dark gray bases which show through the cream color 

One in cream phase is a parous female (USNM No. 307833) whose 
adulthood is further attested by complete ankylosis of the sagittal 
suture posterior to the nasals and the sutures about the interparietal, 
presence of all upper premolars fully erupted, and substantial wear 
on the cheek teeth. The other two in cream phase, however, are 
immatures (cheek teeth without wear; upper premolars deciduous; 
sagittal suture open; interparietal outlined by open sutures) taken 
July 16 and 25 and are evidently molting to gray. The ten other 
June and July immatures do not have quite all of the above indica- 
tions of immaturity, and possess the black or gray dorsal pelage that 
is the ordinary condition or phase. Nothing appears to distinguish 
the pelage of these 10 from that of ordinary adults. 

This subspecies is named for Dr. Boonsong Lekagul of Bangkok, 
Thailand, in recognition of his efforts to arouse in the Thai people 
an interest in increasing knowledge of the flora and fauna of Thai- 
land through establishing, supporting, and using a national museum 
of natural history. 












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Callosciurus caniceps (Gray) 

Definition. — Callosciurus caniceps is a species that includes sub- 
species concolor and adangensis that are extraterritorial, being entirely 
south of the Isthmus of Kra, and himaculatus, domelicus, and caniceps 
in the Indochinese Subregion. See also the synonymy of each of 
these. The range of the species caniceps is shown in Figure 15. 

Diagnosis. — Within the Indochinese Subregion this species is dis- 
tinguished from all others by the combined characters of cool gray 
ventral pelage and a short, abruptly marked off, intensely black tip 
to the tail. 

Relationships to other species. — The range of this tree squirrel is 
from Malaya north to the Salween River in Burma and to 19° 25' N. 
latitude (Doi Chieng Dao) in northern Thailand by our records. It 
is also distributed eastward across Thailand to the Mekong, but 
curiously we find no record of it in eastern Thailand south of about 
14° north latitude (Ban Kabin Buri) nor any record from Cambodia. 

Throughout its range species caniceps is sympatric either with 
species flavimanus or with finlaysoni. It is usually more of a lowland 
species than is flavimanus in the regions in which these two are sym- 
patric, but where the two tend to occur together in generally low 
country caniceps is the larger animal. The species finlaysoni occurs 
with caniceps in the lowlands throughout central Thailand, and here 
also caniceps seems always to be the larger. See the accounts of 
species phayrei and inornaius for relationships of species caniceps 
to these. 

Table 9 provides some body and skull dimensions of 23 of the 
type specimens of named forms that belong in this species. 

Callosciurus caniceps concolor Blyth [Extraterritorial] 

Sciurus concolor Blyth, 1855, Jour. Asiatic Soc. Bengal, 24, p. 474. 
Sciurus lancavensis Miller, 1903, Smithsonian Misc. Coll., 45, p. 16. 
Callosciurus erubescens Cabrera, 1917, Bol. Real. Soc. Espan. Hist. Nat., 17, 

p. 518. 
Callosciurus concolor telibius Thomas and Robinson, 1921, Ann. Mag. Nat. 

Hist. (ser. 9), 7, p. 121. 

Types. — Sciurus concolor, "From the vicinity of Malacca," In- 
dian Museum (not seen); erubescens, MNCN No. 1.669 adult ob- 
tained in Selangor in May or June of 1900 by J. Waterstradt (not 
seen) ; lancavensis, USNM No. 104390, young male taken on Lankawi 
Is. December 1, 1899, by W. L. Abbott; telibius, BM No., 
old male taken on Pulau Telibun, Trang, S.W. Siam, Jan. 2, 1917. 


Material examined, from Malaya. — Straits Settlements [Malacca] 
(MCZ), one, (UMMZ), three, (BM), one; Kuala Lumpur (USNM), 
one, (BM), one; Gin ting Bidai, 2300 feet, Selangor (BM), one; 
"Pahang River" (BM), one; Semangko Pass, Selangor-Pahang 
boundary (BM), one; "Panjum" Kuala Lipis, Pahang (BM), two; 
Telom River, 400 feet (BM), four; Gunong Ijau, 4700 feet, Perak 
(BM), one; Maxwell's Hill, 3600 feet, Perak (BM), one; Temengeh 
[also Temongoh and Tumangoh], 400 feet, upper Perak (BM), one; 
Ulu Selama [also Ulu Selamar], Perak (BM), five; Sungei Lebeh, 
Kelantan (NR), one; Kroh, 1100 feet, upper Perak (BM), nine; 
Gunong Gajah, Kedah (BM), one. 

Material examined, from lower peninsular Thailand.— Bangnara 
[Bang Nara = Narathiwat] (USNM), 16, (MCZ), one, (AMNH), one, 
(CNHM), one, (UMMZ), one; Lankawi Island (BM), 12, (USNM), 
four; Bukit Patani (USNM), two; Patani (BM), two; Biserat, Jalor 
(BM), five; Ban Sai Kau [Nawngchik District] (BM), two; Singora 
[Songkhla] (USNM), two; "Pak Bayoon" (USNM), two; Nakon 
Sritamarat [Nakhon Si Thammarat] (USNM), eight, (ANSP), one, 
(CNHM), one; Pulo Telibun (BM), three. 

Original description. — "Lower-parts dull ash-colour: the rest griz- 
zled throughout with black and dull ruddy-ferruginous; the latter 
somewhat brighter on the middle of the back, croup, and upon the 
tail, which last is conspicuously ringed with black and dull ferrugi- 
nous, and has a black tip mingled with hoary- white." 

Diagnosis. — The above description fits quite well the material in 
the United States National Museum. The diagnostic characters are 
(1) the reddish suffusion of color occurring on the middorsum but 
not on the sides of the neck and body, (2) the actual absence of a 
sharply marked off, quite black tip of the tail, and (3) the cool sil- 
very gray color of the venter. 

Discussion. — It may be seen that material from the northwest 
edge of the range of this subspecies from Pulo Lankawi (named form 
lancavensis) and Pulo Telibun (named form telibius) and the north- 
east edge at Nakon Si Thammarat is intermediate between concolor 
and bimaculatus because it has the sharply marked-off, black tip of 
the tail, but it is listed here for convenience because it is otherwise 
like concolor. See Figure 15. 

Habits. — Near Kuala Lumpur Harrison and Traub (1950, p. 339) 
tabulate data showing that Callosciurus caniceps was frequently 
taken in the forest, was observed in numbers in the town of Kuala 


Lumpur, and was observed in abandoned clearings on the edges of 
country settlements, but that it was not found in the mountain forest 
about 5000 and 6000 feet of elevation on Gunong Brinchang near 
Cameron Highlands. On the other hand Callosciurus flavimanus was 
not taken in the lowland forest about Kuala Lumpur, nor in the 
town, nor in abandoned clearings but was observed in the above 
mountain forest. They also report of caniceps, "This squirrel is very 
common in secondary forest and in association with man. It builds 
nests in trees and feeds on fruit. Thirty-seven specimens were ex- 
amined from the three forest sites," Later Harrison (1954, p. 161) 
reported further on its food, "Although this squirrel is abundant in 
waste land only a few shot specimens have been examined [for food]. 
The stomachs contained fruit and other vegetable matter, and one 
out of three contained a proportion of insects." 

Robinson (in Bonhote, 1903, p. 21) comments from his own and 
Annandale's field notes: "This species is emphatically the village 
squirrel of the Patani States, and it is very exceptionable to find it 
otherwise than in the immediate proximity of dwellings. It is ex- 
ceedingly abundant in the cocoanut groves and orchards, and com- 
mits great ravages among the fruit, being particularly destructive to 
the jack fruit or nangka (Artocarpus integrifolia). It is commonly 
seen on the trees in the early morning, up till about 9 a.m., and after 
about 4 p.m., and in the heat of the day remains hidden in the crowns 
of the palms, where it also forms nests similar to the drey of the Brit- 
ish species. In South Perak, if it occurs, it must be very rare, and 
we never saw a specimen, but in the neighbourhood of Kuala Lumpur 
it, or a closely allied species, is fairly abundant. An entirely black 
variety was seen at Biserat on several occasions." 

Audy and Harrison (1953, p. 10) comment "These . . . squirrels 
are abundant in the understory layer of forest and secondary forest, 
and come down to ground level freely. They range into scrub and cul- 
tivated country, and are well known in the gardens of town houses." 

Callosciurus caniceps adangensis (Miller) [Extraterritorial] 

Sciurus adangensis Miller, 1903, Smithsonian Misc. Coll., 45, p. 17. 

Sciums concolor terutavensis Thomas and Wroughton, 1909, Ann. Mag. Nat. 

Hist. (ser. 8), 4, p. 535. 
Callosciurus moheius Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. 

(ser. 9), 7, p. 122. 
Callosciurus moheius mohillius Thomas and Robinson, 1921, Ann. Mag. Nat. 

Hist. (ser. 9), 7, p. 122. 


Types. — Sciurus adangensis, USNM No. 104389, a young male 
taken on Adang Island, Butang Islands, southwestern Thailand, on 
December 14, 1899, by W. L. Abbott; terutavensis, BM No., 
an old male taken from Telok Udang, Teratau Island, Thailand, 
March 7, 1909, by H. C. Robinson; moheius, BM No., a 
female taken on Pulau Mohea (northern half), 7° 14' N. latitude 
about 30 miles east of Siamese coast, February 2, 1919; mohillius, 
BM No., a male taken from the southern half of Pulau 
Mohea February 3, 1919. 

Material examined, all from islands off southwestern peninsular 
Thailand.— Pulo Terutau (BM), nine, (USNM), eight; Terutau Is- 
land [Pulo Terutaul (BM), nine; Pulo Adang, Butang Islands 
(USNM), two; northern part, Mohea Island (BM), three; southern 
part Mohea Island (BM), three. 

Discussion. — This subspecies consists as presently known, of in- 
sular populations off the most southwestern coast of Thailand in the 
area of intergradation between concolor and bimaculatus. Their pel- 
age characters include the sharply demarked tail tip of bimaculatus 
and the unornamented side pelage and venter of concolor. Although 
they vary somewhat among themselves in pelage color from one is- 
land to another, the diagnostic bright buffy midline of the ventral 
aspect of the tail links them together and distinguishes them from 
both concolor and bimaculatus. 

Callosciurus caniceps bimaculatus (Temminck) 

Sciurus bimaculatus Temminck, 1853, Equisses Zoologiques sur la cote de 

Guine, p. 251. 
Sciurus epomophorus Bonhote, 1901, Ann. Mag. Nat. Hist. (ser. 7), 7, p. 272. 
Sciurus davisoni Bonhote, 1901, Ann. Mag. Nat. Hist. (ser. 7), 7, p. 273. 
Sciurus sullivanus Miller, 1903, Smithsonian Misc. Coll., 45, p. 17. 
Sciurus matthaeus Miller, 1903, Smithsonian Misc. Coll., 45, p. 19. 
Sciurus lucas Miller, 1903, Smithsonian Misc. Coll., 45, p. 20. 
Sciurus epomophorus milleri Robinson and Wroughton, 1911, Jour. Federated 

Malay States Mus., 4, p. 233. 
Sciurus concolor samuiensis Robinson and Kloss, 1914, Ann. Mag. Nat. Hist. 

(ser. 8), 13, p. 226. 
Callosciurus epomophorus nakanus Thomas and Robinson, 1921, Ann. Mag. 

Nat. Hist. (ser. 9), 7, p. 120. 
Callosciurus epomophorus mapravis Thomas and Robinson, 1921, Ann. Mag. 

Nat. Hist. (ser. 9), 7, p. 120. 
Callosciurus epomophorus panjius Thomas and Robinson, 1921, Ann. Mag. 

Nat. Hist. (ser. 9), 7, p. 119. 


Callosciurtis epomophorus panjioli Thomas and Robinson, 1921, Ann. Mag. 
Nat. Hist. (ser. 9), 7, p. 120. 

Callosciurus epomophorus tacopius Thomas and Robinson, 1921, Ann. Mag. 
Nat. Hist. (ser. 9), 7, p. 121. 

Callosciurus epomophorus pipidonis Thomas and Robinson, 1921, Ann. Mag. 
Nat. Hist. (ser. 9), 7, p. 121. 

Callosciurtis epomophorus tabaudius Thomas, 1922, Jour. Bombay Nat. Hist. 
Soc, 28, p. 1067. 

Callosciurus epomophorus hastilis Thomas, 1923, Jour. Bombay Nat. Hist. 
Soc, 29, p. 377. 

Types. — Sciurus bimaculatus, RNH No. 13363, an adult male 
from the peninsula of Malacca; epomophorus, BM No., 
old female, from Salanga Island, Thailand, collected March 4, 1879, 
by J. Darling; davisoni, BM No., young female, from Ban- 
kasun, Tenasserim, Burma, collected June 20, 1877, by W. Davison; 
sullivanus, USNM No. 104377, old female, from Sullivan Island, 
Burma, collected February 1, 1900, by W. L. Abbott; matthaeus, 
USNM No. 111920, adult female, from St. Matthew Island, Burma, 
collected December 11, 1900, by W. L. Abbott; lucas, USNM No. 
104385, adult female, from St. Luke Island, Burma, collected Janu- 
ary 20, 1900, by W. L. Abbott; milleri, BM No., young male, 
from Trong, Thailand, collected February 20, 1896, by W. L. Ab- 
bott; samuiensis, BM No., adult male, from Samui Island, 
Thailand, collected May 12, 1913, by H. C. Robinson and E. Sei- 
mund; nakanus, BM No., old male, from Naka Island, 
Thailand, collected February 4, 1918, by H. C. Robinson and C. B. 
Kloss; mapravis, BM No., adult female, from Maprau 
Island, Thailand, collected February 10, 1918, by H. C. Robinson 
and C. B. Kloss; panjius, BM No., old male, from Pan- 
jang Island, Thailand, collected January 20, 1918, by H. C. Robin- 
son and C. B. Kloss; panjioli, BM No., adult male, from 
Panjang Anak Island, Thailand, collected January 29, 1918, by 
H. C. Robinson and C. B. Kloss; tacopius, BM No., old 
male, from Takopah Island, Thailand, collected February 16, 1919, 
by H. C. Robinson and C. B. Kloss; pipidonis, BM No., 
adult female from Pipidon Island, Thailand, collected February 3, 
1919, by H. C. Robinson and C. B. Kloss; tabaudius, BM No., an adult female taken on Tavoy Island, Mergui Archi- 
pelago, Burma, October 21, 1921, by C. Primrose; hastilis, BM No., adult male, from Hastings Island, Burma, collected Oc- 
tober 21, 1921, by C. Primrose. 


Material examined, from mainland of peninsular Thailand. — Tha 
Lo [Ban Tha Lo], Bandon (USNM), three; Bandon [Amphoe Ban 
Don=Ban Makham Tia] (USNM), two; Trang (USNM), eight, 
(ANSP), one, (CNHM), one; Ban Ta [Tha] Yai (USNM), one; Kao 
[Khao] Luang (USNM), one, (CNHM), one; "Waterfall," Trang 
(USNM), two; Kao [Khao] Chong (BM), three, (USNM), one; Kao 
[Khao] Soi Dao (USNM), two; Poonga [Phangnga] (MCZ), one, 
(BM), two; Tang Pra [Laem Ao Kham], Takuatung (BM), four; 
Taptien [Tapting], Trang (BM), five; Chong, Trong (BM), three; 
Gap Patalong, Trang (BM), one; "Lem Ma," Trang (BM), three; 
Nam Chuh, Pak Chan (BM), one; "Renong River" [Ranong] (BM), 
two; Klong [Khlong] Wang Hip, Tung Sawng [Thung Song] (BM), 
one; Mamoh, Pakchan (BM), two; Torsan [Ban Tha San], Chun- 
tawn [M. Chum Pon] (BM), one; Nong Kok, Grabi [Ban Nong Kok, 
Krabi] (BM), one. 

Material examined, from Siamese islands west of peninsula. — 
Tongka [Koh Phuket] (BM), eight; Telok Palas [Ban Ao To Nong, 
Koh Phuket] (BM), two; DeLisle, or Piam, Island (BM), four; 
"Pasir Raja Is." (BM), six; Pipidon Island (BM), two; Maprau Is- 
land (BM), four; Panjang Island (BM), two; Panjang Anak [Island] 
(BM), three; Naka Island (BM), three; Takopah Island (BM), 
three; Pulo Sarik, Tongka [Ko Sire, near Ko Phuket] (BM), five. 

Material examined, from Siamese islands east of peninsula. — Koh 
Samui [Ko Samui] (USNM), four. 

Material examined, from Tenasserim mainland, Burma. — Bok Pyin 
(USNM), one; Maliwun (USNM), two; Victoria Point (USNM), 
three, (BM), one; Sungei Balik (USNM), one; Tanjong Badak 
(USNM), two; Champang (USNM), one; Boyce's Point (USNM), 
one; Bankasoon (BM), 14 topotypes; Taroar [Tagoot], Malay Pen- 
insula (BM), one; "Kussoom," N.W. peninsula, south of Tenasserim 
(BM), one; Tenasserim Town (BM), one. 

Material examined, from Mergui Archipelago, Burma. — St. Mat- 
thew's Island (USNM), four; St. Luke's Island (USNM), one; Has- 
tings Island (BM), three; Sullivan Island (BM), two, (USNM), five; 
Clara Island (USNM), one; James Island (USNM), three; Tavoy 
Island (BM), two; King Island (BM), six; "Nathay Mine" (BM), 
six; Kisseraing Island, 100 feet (BM), three; "Sir John Malcolm 
Island," 100 feet (BM), two. 

Original description. — Temminck's species bimaculatus clearly be- 
longs here, for his description of it nicely provides the pelage charac- 
ters diagnostic for this subspecies: "The upper parts of the head. 


neck, all the back and the tail as far as the tip bear a . . . pelage regu- 
larly ringed with ashy and black . . . the sides of the neck, the upper 
part of the legs and flanks are of the red of rust; the lower parts are 
of a whitish gray." 

Another very early specimen erroneously referred by Horsfield 
(1824, no. 7, p. 8) to Sciurus affinis Raffles, 1822, is sometimes {e.g., 
Ellerman, 1940, p. 355) treated as if it were the type of another 
affinis, even though Sciurus affinis Raffles is at the same time recog- 
nized as valid for a species of Ratufa. The description by Horsfield, 
and his specimen do clearly belong to what is now another genus, 
Callosciurus, and in the present species and subspecies, but Sciurus 
affinis Horsfield, 1824, is a primary homonym of Sciurus affinis 
Raffles, 1822, and is permanently rejected (Mayr, Linsley and Usin- 
ger, 1953, p. 312), 

Diagnosis. — The present subspecies is characterized by being gray 
where concolor is reddish (i.e., on the middorsum) and reddish on the 
sides where concolor is gray. There is considerable variation in in- 
tensity of both the gray and the red, and some of it is geographic, 
but these differences adequately distinguish the present subspecies 
from the localities presently indicated. Callosciurus caniceps bi- 
maculatus also has a sharply black-tipped tail, and as mentioned 
above, this also occurs on material which is geographically intergrade 
between concolor and bimaculatus. The type specimen of bima^u- 
latu^ does also have a white tip distal to the black tip, but in the 
opinion of the one of us (Tate) who examined it, the white tip is an 
individual aberration. 

The reddish infusion of the color on the sides which characterizes 
bimaculatus extends over the side of the neck posterior to the ear and 
along the side of the body, posteriorly extending ventrally onto the 
groin. It may or may not be continuous from the neck across the 
shoulder to the side of the body and from the side onto the thigh. 
In areas where the red side color is most intense, it tends to be so 
continuous; in areas where the red color pales, the side of the neck 
and groin retain the most color almost as a large spot in each place. 
Koh Phuket is a focal geographic area for intensification of the red. 

Pelage color. — The venter is generally gray with an overlay of 
whitish hair tips, and a darker gray midline of agouti pelage, but in 
areas where the sides are intensely red, there may be a general suf- 
fusion of reddish invading the ventral pelage. 

Habits. — One highly qualified observer collecting this squirrel in 
most southern, peninsular Burma comments: "Very plentiful every- 


where, both around habitations and in the thickest forest. A thick- 
set rather clumsy looking squirrel. Wherever there are plantations 
this squirrel destroys large numbers of cocoanuts, by drilling a cir- 
cular hole in the side and extracting the contents. Weight. 10-14 
ozs." (G. C. Shortridge in Wroughton, 1915b, p. 713.) 

Discussion. — In the collections of the named forms of the main- 
land north of subspecies concolor that are available in the United 
States National Museum, we laid out 16 specimens of "davisoni" from 
the Isthmus of Kra near the tenth parallel of latitude north (see 
Figure 15), and 14 specimens of "milleri" from Trong, represented 
in Figure 15 by the southernmost all-black dots. We could find no 
consistent difference between these either in color or size, both lots 
proving quite variable and the number of individuals showing inter- 
mediacy in any particular character often exceeding the number that 
show a discrete difference. 

Although St. Matthew's Island and Sullivan Island of the Mergui 
Archipelago are both outside the nine meter depth line (Fourth Sur- 
vey of India, U. S. Army Map Service, 1944, sheet N. C. 47) the 
named forms from them, matthaeus and sullivanus, are indistinguish- 
able from each other by comparison of the original hypodigms (5 and 
6 specimens respectively) and are indistinguishable from the series 
of thirty specimens of bimaculatus from the adjacent mainland. St. 
Luke's Island and Hastings Island are satellite islands of St. Mat- 
thew's Island, each less than a mile from it and connected to it by 
shallows less than nine meters deep. The original hypodigm of two 
specimens of the named form from St. Luke's I., lucas, are distin- 
guishably darker than all of the mainland sample of bimaculatiLS, 
but not the St. Matthew's I. sample. In lumping the Hastings I. 
form, hastilis, with bimaculatus {= davisoni) , we follow Ellerman and 
Morrison-Scott (1951, p. 486). 

Callosciurus caniceps casensis (Miller) [Extraterritorial] 

Sciurus casensis Miller, 1903, Smithsonian Misc. Coll., 45, p. 19. 

Type. — USNM No. 104370, a young female taken on Chance 
Island, Thailand, December 28, 1899, by W. L. Abbott. 

Material examined. — Chance Island, off west coast of peninsular 
Thailand (USNM), four. 

The original hypodigm of five specimens, all we have seen, vary 
little among themselves. They have the ventral side of the tail strik- 
ingly paler than those of any specimens of geographically nearby 


forms of caniceps and grayer than the somewhat yellowish ones of 
altinsularis, which are from High Island 110 miles to the north. 

Callosciurus caniceps altinsularis (Miller) 

Sciurus altinsularis Miller, 1903, Smithsonian Misc. Coll., 45, p. 21. 

Type.— VSNM No. 111975, an old female taken on High Island, 
Burma, December 31, 1900, by W. L. Abbott. 

Material examined. — High Island, Mergui Archipelago, Burma 
(USNM), four. 

The four specimens and the type laid out together, vary hardly 
at all among themselves, and do not differ ventrally from several of 
the himaculatus in the United States National Museum, but the dor- 
sal pelage as a whole is slightly but distinctly paler than any of the 
himaculatiLs and paler than any of the nearby insular forms except 
casensis (which, however, is much more brilliantly colored). 

Callosciurus caniceps fallax (Robinson and Kloss) 

Sciurus concolor fallax Robinson and Kloss, 1914, Ann. Mag. Nat. Hist. (ser. 8), 
13„ p. 225. 

Type. — BM No., an adult male taken on Pennan (or 
Pangnan) Island, Thailand, 9° 45' N. latitude off east coast, on 
May 30, 1913. 

Material examined. — Koh Pangan, Thailand (USNM), four. 

The material examined is consistent within the series and distin- 
guishable from bimaculatus by having paler tails (quite like those of 
casensis). It differs from casensis by quite dull pelage, the ruddiness 
of the sides being barely noticeable, and differs from altinsularis by 
much darker pelage on both dorsum and venter, and very much 
greater size. 

It seems a bit odd that the principal distinguishing character of 
fallax should be one shared only with two insular forms on the far 
side of the Malay peninsula, casensis 140 miles west and very slightly 
south, and altinsularis about 140 miles northwest. 

Callosciurus caniceps domelicus (Miller) 

Sciurus domelicus Miller, 1903, Smithsonian Misc. Coll., 45, p. 18. 
Sciurus bentincanu^ Miller, 1903, Smithsonian Misc. Coll., 45, p. 19. 

Types. — Sciurus domelicus, USNM No. 104381, an adult female 
taken on Domel Island, Mergui Archipelago, February 24, 1900, by 


W. L. Abbott; bentincanus, USNM No. 104383, an adult female 
taken on Bentinck Island, Mergui Archipelago, March 11, 1900, by 
W. L. Abbott. 

Material examined. — Domel Island (USNM), four; Bentinck's 
Island (USNM), three; Kisseriang Island (USNM), one. 

The sample of five specimens from Domel Island representing 
this form is a darker gray ventrally than any of the material of hi- 
maculatus examined (by Moore). Since the Bentincks Island mate- 
rial is also rather dark dorsally and intensely red laterally like the 
material from Domel Island, and is separated geographically by 
Domel Island from the mainland range of himaculatus, it is here in- 
cluded in the subspecies domelicus. The subspecies domelicus also 
occurs on Kisseriang Island which lies between Domel Island and 
the mainland. 

Callosciurus caniceps caniceps (Gray) 

Sciurus caniceps Gray, 1842, Ann. Mag. Nat. Hist. (ser. 1), 10, p. 263. 

Sciurus chrysonotus Blyth, 1847, Jour. Asiatic Soc. Bengal, 16, p. 873. 

Sciurus epomophorus inexpectatus Kloss, 1916, Jour. Nat. Hist. Soc. Siam, 2, 
p. 178. 

Sciurus helgei Gyldenstolpe, 1917, Kungl. Svensk. Vet. Handl., 57, no. 2, p. 34. 

Sciurus caniceps hehus Shamel, 1930, Jour. Mammal., 11, no. 1, p. 72. 

Sciurus epomophorus fluminalis Robinson and Wroughton, 1911, Jour. Fed- 
erated Malay States Mus., 4, nos. 3 and 4, p. 233. 

Types. — Sciurus caniceps, BM No. 213a (41.1817), an adult male 
from Tenasserim; chrysonotus (not seen), from Tenasserim Valley; 
inexpectatus, USNM No. 221557, a young female (deciduous upper 
fourth premolar still present) from Koh Lak, Pran, latitude 11° 45' N., 
southwest Siam, collected November 15, 1916 by C. B. Kloss; helgei, 
NR No. 71, a young male from Koh Lak, Siam, collected Novem- 
ber 29, 1914, by N. Gyldenstolpe; fluminalis, BM No., an 
adult male taken at the Meping Rapids, Siam, August 8, 1907, by 
T. H. Lyle. 

Material examined, all from Koh Tau, Thailand (USNM), 12. 

Material examined, from the mainland of Thailand. — Khao Luang 
(ANSP), five, (CNHM), one; Hat Sanuk, near Koh Lak, Rajburi 
(BM), six; Pran [Prachuap Khiri Khan] (USNM), five; Koh Lak 
[Prachuap Khiri Khan] (CNHM), one, (USNM), five, (NR), one, 
(BM), eight; Sam Roi Gop [Sathani Sam Roi Yot] (USNM), four; 
Kwe Koi [Mae nam Khwae Noi] (USNM), one; Muang Kan Buri 


[Kanchanaburi] (CNHM), one, (USNM), two; Ban Pong, Rajburi 
(USNM), one; "Nong Mong, Muang Krabin" [Ban Kabin Buri] 
(USNM), two; "Lam Ton Lang, Krabin" [Ban Kabin Buri] (USNM), 
two; Krabin [Ban Kabin Buri] (BM), three; Kao Lem [Khao Laem] 
(USNM), two; Pak Jong [Ban Pak Chong] (AMNH), four, (CNHM), 
one, (USNM), two; "Lam Klong Lang, Pak Chong" [Ban Pak Chong] 
(USNM), one; Muek Lek [Ban Muak Lek] (CNHM), one; Lat Bua 
Kao [Ban Lat Bua Khao] (USNM), one; Siken, near Korat [Ban Si 
Khiu] (USNM), one; Chainat [Muang Chainat] (BM), one; Pak- 
nampo [Ban Pak Nam Pho] (MCZ), one, (BM), two; Nakon Sawan 
[Nakhon Sawan] (BM), two; Bung Borapet [Boraphet] (USNM), 
one; Um Pan [Ban Um Phang=Ban Le Kathe] (AMNH), two, (BM), 
two; Longlung [near Ban Nong Pla Lai] (BM), one; Wung Pratart 
Farm, Kampengpet Prov. (CNHM), three; Klong Klung [Ban 
Khlong Khlung] (CNHM), five; Kampengpet [Changwat Kam- 
phaeng Phet] (AMNH), one, (CNHM), two, (BM), one; Sawan 
Kaloki [SawankhaIok=Ban Wang Mai Khon] (BM), three; Pak Koh 
(NR), two; Ban Hue Hom (NR), one; Mee Tan [Ban Mae Tan Nua] 
(BM), two; Mt. Chieng Dao (USNM), two, (AMNH), three; 
"Watpa" (ANSP), one; "Ubol Chanumon" (ANSP), one; "Nong 
Dom" (UMMZ), three; "Me Ping River" (AMNH), one; (BM), 
four; "Ban Kon" (NR), one; "Kao Phlyng" (NR), one; Phan Mt., 
Sakon Nakhon (USNM), two; Lomlo Mt., Ban Maeo (USNM), 17; 
"Phak Khinak Mt.," Dan Sai (USNM), one; "Nam Lang Mt.," 
Ban Khok (USNM), one; Kowjeen, Pak Tho, Rajburi (USNM), 
two; "Pukongchai," Korat (USNM), one; Hin Laem, Tra Khanun, 
Kanchanaburi (USNM), 16; Ban Klua Klang, Prachuap Kiri Khan 
(USNM), three; "Ban Sop Luak," Chiang Saen Kao, Chiang Rai 
(USNM), one; "Mushi Tar Shang," Korat (USNM), one; "Ban Hua 
Thanon," Klong Klung (USNM), six; Kowkat, Paknampho (USNM), 
16; Kowkob, Paknampho (USNM), two; "Moung Wat Sy Tie," 
Nakorn Sawan (USNM), one; "Sretahn, Wung Sapueng," Loei 
(USNM), one. 

Material examined, from Burma. — Tavoy (BM), two; "Shan 
Mepa," Amherst (BM), one; "Tikotaw" Amherst (BM), one; Lakya 
(AMNH), two; Kawlichuang (AMNH), one; Lampha (AMNH), 
one, (BM), three; Moulmein (BM), three; Kawkareik (AMNH), one, 
(BM), one; Myawadi (BM), one; "Lothorgu," Myawadi (BM), one; 
"Thoungyin River, 1500 feet" (BM), three; "Thoungyin above 
Myawadi" (BM), three. 


Original description. — Gray's type description of caniceps is very 
bare: "Pale gray, grisled; back yellowish, beneath paler gray; tail 
long, gray, black varied, ringed, hair with three broad black bands." 

Pelage color. — Dorsal pelage color of American Museum Tenas- 
serim material is about Orange Rufus (II) and that of Doi Chieng 
Dao about Ochraceous Orange (XIV). Ventral pelage color is an 
agouti of about Deep Gull Gray (LI 1 1) in this Tenasserim and mid- 
dle Siam material, and about Light Gull Gray in the northernmost. 
Every American Museum specimen has a darker agouti, longitudinal 
midventral stripe about four to six millimeters wide. The throat, 
chin and under sides of legs are like the general color of the venter. 
The tail hairs have five black bands besides the black tips. The tips 
of the tails are all abruptly black. Dorsally the tail and all legs are 
agouti of about Mouse Gray (LI). The dorsal pelage of the feet, 
snout and ear tips is notably lighter and corresponds closely to the 
ventral pelage color. The bright orange color covers the back and 
sides, fading into agouti gray as it approaches the venter, on the 
crown, and on the proximal one- tenth of the tail. 

Habits.- — Gyldenstolpe (1914, p. 11) remarks interestingly, "In 
Siam this species was only common in the bamboo-forests in the 
North and seemed to live in rather high altitudes. In the bamboo- 
forests on the Korat plateau in Eastern Siam it was never observed." 

Discussion. — Although the named form inexpectatus is distin- 
guishable when one compares series of study specimens in a museum, 
it seems to occupy but a small geographic area between the ranges 
of subspecies bimaculatus and subspecies caniceps, the rainshadow 
area of peninsular Thailand north of about 11° North latitude. 
Since its observed characters in the material examined are but tran- 
sitional between those of bimaculatus and caniceps, it is here regarded 
as intermediate between those two subspecies and not fully belong- 
ing to either, but for convenience the records of it are lumped with 
caniceps. The named form helvus from Koh Tao is rather distinct, 
but the real interest which attaches to the Koh Tao material is that 
it shows relationship as close to caniceps as to the geographically 
nearer bimaculatus, and like the sample of "inexpectatus" is actually 
intermediate. Inclusion of it with subspecies caniceps emphasizes 
the interest which attaches to its close relationship to caniceps across 
a much greater water gap. It is interesting, too, that the sample 
from Koh Phangan fallax, which lies between the Koh Tao popula- 
tion and the bimaculatus of the nearest mainland is good bimaculatus, 
but shares with the Koh Tao sample a character which differs from 
mainland bimaculatus (pale underside of tail). 



It certainly seems worth mentioning here that in the American 
Museum of Natural History material the roughly topotypical cani- 
ceps specimens from the heavy rainfall area of upper Tenasserim are 
not darker colored than ones from the much drier, rainshadow area 
of Siam almost directly east (localities Kawkereik, Lampha, Kaw- 
lichaung, and Lakya in Burma, versus Um Pang, Me Ping, and Kam- 
pengpet, Siam). The northernmost material that we have seen for 
caniceps, from Doi Chieng Dao, is notably lighter in both dorsal and 
ventral pelage, and is also slightly larger than this more southern 
American Museum material. If these differences could be shown to 
be consistent over a considerable geographic area, the name flumi- 
nalis Robinson and Wroughton would be available for the subspecies. 

The orange-colored dorsal pelage of Callosciurus c. caniceps has 
been reported to change to agouti gray color seasonally. The follow- 
ing records from examination of museum specimens in London (by 
Tate) and Chicago, Boston, and New York (by Moore) provide some 
evidence of such seasonal change: January, 12 orange, no gray; Feb- 
ruary, four orange, no gray; March, 12 orange, one changing, one 
gray; April, one orange, two changing, no gray; May, no data; June, 
one orange, three changing, nine gray; July, no orange, one changing, 
two gray; August, no data; September, no orange, one gray; October, 
one orange, two changing, no gray; November, three orange, one 
gray; December, two orange, one changing, no gray. It thus appears 
that in the November through April or May dry season the pelage 
is more often orange, but during the June through October season 
of rains, the dorsal pelage is more often gray. 

Some of the specimens that we have recorded above as "chang- 
ing" may possibly be only intermediate in some other sense. How- 
ever, one (CNHM No. 47332) in April was definitely molting the 
orange and replacing it with gray, and one December specimen 
(AMNH No. 54692) seems to be changing from gray to orange with- 
out showing a molt line, by the simultaneous appearance of orange 
in large patches throughout the length of the back. In this latter 
specimen the guard hairs of the remaining patches of agouti gray 
pelage seem under magnification quite as fresh as those of the in- 
cipiently orange blotches. Furthermore, the guard hairs of the incipi- 
ently orange blotches are agouti and only warmly colored on the two 
light bands (more deeply colored on the proximal one), and more of 
the orange color seems attributable to the thinner under hairs. How- 
ever, in a fully orange specimen of January 19th (AMNH No. 54707) 









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from nearby, the guard hairs are not agouti but are entirely orange 
excepting the thin, short, black tip and a short paler base, and there 
seems to be very little pelage wear (breakage of these guard hairs). 
In a fully orange March 3rd specimen, the pelage is as in the preced- 
ing specimen but shows wear (has the tips of a good many guard 
hairs broken off). These few superficial observations suggest that at 
the end of the rainy season, orange color may replace the black and 
white bands on the agouti guard hairs rather than orange guard hairs 
replacing black and white banded guard hairs by molt. Thus it may 
be that this tropical tree squirrel molts but once annually while 
accomplishing change in color twice a year, although it seems well 
established that the well-studied north temperate tree squirrels Tami- 
asciurus hudsonicus, Sciurus vulgaris, and Sciurus carolinensis are 
known to molt twice a year (each spring and fall). 

Callosciurus phayrei (Blyth) 

Sciurus phayrei Blyth, 1855, Jour. Asiatic Soc. Bengal, 24, p. 472. 
Sciurus blanfordi Blyth, 1862, Jour. Asiatic Soc. Bengal, 31, p. 333. 
Callosciurus griseimanus heinrichi Tate, 1954, American Mus. Novitates, no. 
1676, p. 1. 

Types. — Sciurus phayrei, BM No., an adult male taken 
at Martaban, Burma, by Major Phayre; blanfordi BM No. 63.5.- 
9.9, an immature taken at Mt. Ava about 20 mi. S.W. of Mandalay; 
heinrichi AMNH No. 163466, an adult female taken at Maymyo, 
Mandalay, Burma, at 800 meters elevation on December 6, 1937, by 
Gerd Heinrich. 

Material examined, all from Burma, south to north. — Martaban 
(BM), one cotype; Thatone [Thaton] (BM), three; Pahpoon [Papun] 
northern Tenasserim (BM), two; "Bantam, Kiu Pang Valley," Sal- 
win Distr. (USNM), one; five miles east of Toungoo, 100 feet 
(CNHM), two; 40 miles north of Toungoo, 500 feet (BM), four; 
"Ta-ho, Kareni" (USNM), one; Mt. Ava (BM), one cotype; seven 
miles southeast of Mandalay (USNM), one; Maymyo, 800 meters 
(AMNH), six; Gokteik, 2100 feet (BM), three; Kokkoaing, 40 miles 
N.N.E. of Mandalay, 800 feet (BM), one; Ngapyiniun, 250 feet, 
opposite Kyaukmyaung, on Irrawaddy R. (MNHN), one, (BM), 
six; Pyaunggaung, 2800 feet (BM), three; Hsipaw, 1350 feet (BM), 
two; Se'en, 1400 feet (BM), five, (CNHM), two. 

Table 10 provides some body and skull dimensions of some of the 
type specimens of named forms that belong in species Callosciurus 


Definition. — The species phayrei is accepted here as monotypic 
and is confined to the region between the valley of the Sittang and 
upper Irrawaddy rivers and Salween River as shown in Figure 15. 

Diagnosis. — The species phayrei has the following distinctive color 
characters of the pelage. (1) The rostrum, ears, crown, and side of 
neck are agouti gray like the dorsum. (2) The contrasting color of 
the manus stops abruptly at the wrist. (3) The tail is tipped briefly 
atid rather abruptly with black. (4) The ventral pelage varies from 
a rich orange to a very pale orange but is never red or gray. (5) All 
four feet are yellowish buff to pale orange. (6) Long, yellow bases 
of the ventral tail hairs provide a rich yellow midstripe 12 to 15 mm. 
wide along the under side of the tail. (7) There is usually at least a 
faint blackish band 20 mm. wide along (and within) the lateral mar- 
gins of the ventral pelage. (8) There are five blackish bands on the 
fully grown out tail hairs. 

The above characteristics distinguish species phayrei from the 
subspecies of other species that are geographically closest to phayrei, 
as follows: C. erythraeus sladeni by 1, 2, 3, 4, 5, 6, and 7; C. flavi- 
manus shanicus by 1, 4, 5, 6, and 7; C. flavimanus zimmeensis by 1, 
2, 4, 5, and 7; C . ferrugineous by 1, 2, 3, 4, 5, 6, 7, and 8; C. c. caniceps 
by 4. 5, 6, and 7; C. pygerythrus janetta by 1, 2, 5, 7, and 8. 

Relationships to other species. — This species appears to inhabit the 
rain forest vegetation area from Martaban at the mouth of the Sal- 
ween River north to about Toungoo, and to occupy the strip of trop- 
ical deciduous forest along the escarpment of the Shan Highlands 
from about Toungoo north to at least fifty miles north of Mandalay 
(according to the map of vegetation zones of Burma by de Terra 
(1944, p. 80). On ecological grounds it seems very likely that the 
range of phayrei would extend up the long corridor of rain forest of 
the Salween River valley. Whether it also occurs throughout the 
Shan Highlands seems more doubtful for the Maymyo, Gokteik, 
Pyaungaung, and Se'en localities seem to lie along the southeastern 
margin of an eastward jutting of the tropical deciduous forest into 
the "mixed forest with grassland" of the Shan Highlands (de Terra, 
1944, p. 80), and may not represent habitat typical of the Shan 

Relationships between C. phayrei and C. erythraeus sladeni. — 
While our record of the distribution of phayrei from material ex- 
amined leaves large gaps where its occurrence remains to be ascer- 
tained, its range seems to be confined between two rivers, on the 
south and east by the Salween and on the west by the Irra- 


Fig. 15. Geographic distribution of three species, as determined from mate- 
rial examined: the Tenasserim squirrel, Callosciurus caniceps, circular symbols; the 
Shan Highlands squirrel, Callosciurus phayrei, black squares; and the Laotian 
squirrel, Callosciurus inornatus, triangles. Subspecies of caniceps: half black [extra- 
territorial], concolor; vertically divided [extraterritorial], adangensis; all black, 
bimaculatus; dotted circles, domelicus; and open circles, caniceps. 



waddy. Across the upper Irrawaddy phayrei seems to be ecologically 
replaced by sladeni as the tree squirrel of medium size in the forests 
in the broad fertile valleys and relatively low hills of this interri- , 
parian area of Burma. See map figs. 13 and 15. i 

Besides occupying what is apparently the same ecological niche 
on opposite sides of the Irrawaddy River phayrei and sladeni share 
a number of characteristics of the pelage: (1) The dorsal pelage of 
all four feet is colored like the venter. (2) The dorsal pelage of the 
body and proximal part of the tail are an agouti gray with three light 
bands on many of the guard hairs of the middorsum and with five 
black bands on the hairs of the tail at its midlength. (3) The tail is 
not agouti at its tip. (4) The ventral pelage is orange-red or orange 
but never divided down the midline by an agouti stripe. (5) The 
ears are the same color as the dorsum. 

The pelage characters which distinguish phayrei and sladeni are : 
(1) The specimens of sladeni almost invariably have the rostrum, 
and sometimes the crown back as far as the ears, colored like the 
feet; whereas the rostrum and crown of phayrei are as agouti gray 
as its back. (2) The color of the manus in sladeni extends up onto 
the forearm, meeting the agouti gray abruptly for half the forearm's 
length; whereas in phayrei the manus color terminates abruptly at 
the wrist. (3) In sladeni the tail has a rather long, reddish tip, but 
in phayrei the tail tip is short, fairly abrupt, and black. These three 
characters, in the absence of any known area of intergradation, sug- 
gest something more than subspecific difference between the two. 
Even though the evidence is excellent that the subspecies sladeni 
and haringtoni are conspecific, and although the pelage difference 
between sladeni and haringtoni appears to exceed that between sladeni 
and phayrei, the latter two are not known to intergrade: their differ- 
ences are great enough that they probably would not interbreed if 
brought together naturally, for example, by a shift of the course of 
the Irrawaddy and elimination of the old channel. (Intergradation 
between sladeni and haringtoni is apparently accomplished only 
through another subspecies and interbreeding might not occur at all 
between individuals of these two forms themselves if they could be 
brought together under natural conditions.) 

The Sittang Valley separates the range of C. phayrei from that 
of C. ferrugineus to a great extent, although the specimen of phayrei 
from Ava shows that phayrei has crossed the valley a short distance 
eastward and a specimen each from Gokteik and Lawksawk indicate 
that ferrugineus has crossed the valley westward. It is assumed 


that this condition of Hmited or incipient sympatry has come about 
entirely since the capture of the headwaters of the "Irra-Sittang" 
by the "Chindwin-waddy" {i.e., the removal of the river barrier from 
the Sittang Valley). Their difference in pelage color is virtually total, 
ferrugineus being an all red species with some blackening on the 
feet and tail. This fact and those of sympatry with no evidence of 
intergradation demonstrates that phayrei and ferrugineus are sep- 
arate species. 

Similarities between C. caniceps caniceps and species phayrei. — 
From the material examined, it appears that the subspecies caniceps 
extends northwestward in typical form to the vicinity of the Salween 
River and stops abruptly there; and immediately on the other side 
of the Salween River the species phayrei apparently replaces caniceps 
ecologically. The lack of intergradation between these two forms 
and the niceness of their separation by the existing geographic bar- 
rier of the river are such that no previous student has regarded them 
as conspecific. Nevertheless, the number of evidences of close rela- 
tionship is great: (1) The back, sides and dorsum of the tail of phayrei 
are colored like those of gray phase caniceps. (2) The tail tip in 
phayrei is black like that of caniceps. (3) The head, ears, and rostrum 
of phayrei are gray as in gray phase caniceps. (4) All four feet are 
strikingly light in both forms, lighter than their respective ventral 
pelage. (5) The tail hairs have five black bands additional to black 
tips in both forms, and the guard hairs of the dorsum have two white 
or very pale buffy bands in phayrei as in the gray phase of caniceps. 
(6) The frontal projects forward as a thin process between the nasal 
and the premaxillary on each side, often to a distance equaling least 
width of nasal in many subspecies of Callosciurus; but it does not 
do this in caniceps or in the material representing phayrei in the 
American Museum of Natural History. (7) Subspecies caniceps, 
unlike most mainland forms of large Callosciurus, is strongly pro- 
odont, and the seven Maymyo specimens of phayrei are at least 
somewhat proodont. 

Distinctions between species phayrei and C. caniceps caniceps. — 
(1) The ventral pelage is Light Ochraceous Buff to Ochraceous BufT 
(in the Maymyo series) even approaching Ochraceous Orange on the 
underside of the hind legs, compared to the whitish over pale gray 
of caniceps, and lacks any vestige of the midventral agouti stripe of 
caniceps. (2) All four feet are dorsally Light Ochraceous Buff in 
phayrei (Maymyo series) but about Pale Gull Gray (LIII) in cani- 
ceps (Mt. Chieng Dao series). (3) The ventral aspect of the tail of 


phayrei reveals long, yellow bases of ventral tail hairs so that the 
tail has a Warm Buff midstripe 12 to 15 mm. wide beneath but is 
cool gray dorsally. In caniceps some faint yellow color shows in the 
dorsal aspect but ventrally virtually none. (4) In phayrei there is a 
20 mm. wide blackish band on either side in the ventral pelage about 
60 mm. long. This is very faint in the Maymyo series but also in 
three Gokteik, one Pyaunggaung, and three Se'en specimens in the 
British Museum. The others in the British Museum have the ven- 
tral stripes strongly marked and agree completely in appearance with 
the type. Both variants were thus taken at the Se'en and Pyaung- 
gaung localities and all in May. It is not apparently a seasonal or 
sexual difference, nor is it clearly a geographic variation. (5) The 
ears appear to be notably shorter in the Maymyo series of phayrei 
than in caniceps. (6) Subspecies phayrei lacks the seasonal orange 
dorsal pelage of caniceps. 

C. caniceps caniceps and C. phayrei apparently maintain these 
striking pelage color differences to the opposite banks of the Salween 
River which separates them, and in absence of any indication of in- 
tergradation, and on the degree of difference in pelage characters, 
they are here confidently regarded separate species. 

C. pygerythrus janetta is geographically adjacent to C. phayrei on 
the west and resembles it more closely than do any of the other sub- 
species of C. pygerythrus, excepting possibly the typical subspecies 
pygerythrus (which likewise is only west across the Sittang River 
from C. phayrei and resembles the rainforest population of phayrei 
almost as closely as janetta does the deciduous forest population). 
The close relationship these observations suggest, demands direct 
comparison of janetta with phayrei. 

Just as phayrei shares certain color characters with C. c. caniceps 
as its closest relative to the south or east, phayrei shares some also 
with C. p. janetta revealing it to be the closest relative to the west. 
These latter are: (1) All of the dorsal pelage is a cool agouti gray 
excepting hands and feet and tip of tail. (2) The dorsal pelage of all 
four feet is not agouti but is colored like that of the venter and con- 
trastingly lighter than that of the dorsum. (3) There is an abrupt 
short, black tip to the tail. (4) There is a narrow buffy mid ventral 
stripe along the tail. (5) There is some darkening of the ventral 
pelage in a broad band across the abdomen, tending to fade near 
the midline. The first three of these five characters are shared also 
by C. c. caniceps. 




The important color differences between phayrei and janetta are: 

(1) There are five blackish bands to fully grown out tail hairs in 
phayrei, but four in janetta. (2) There is a flash mark on each hip 
of janetta, none in phayrei. (3) The hind feet of phayrei are nearly 
Warm Buff whereas those of janetta are but Cream Color. (4) The 
ventral pelage of phayrei is contrastingly richer in color than the mid- 
ventral longitudinal tail stripe, but in janetta they are about the 
same, or the venter is the more pale. (5) The side of the neck is 
pale, almost cream color, in janetta, but hardly lighter agouti than 
the nape in phayrei. 

There is an important general size difference between phayrei and 
janetta which is fairly well indicated by hind foot measurements of 
both these forms taken in the field by the same collector, Gerd Hein- 
rich. These are: janetta, 40, 40, 40, 43 mm.; phayrei, 43, 45, 46, 47, 
48, 48 mm. C. phayrei has been taken at Ava west of the Irra- 
Sittang Valley and janetta east of it at Mandalay. Since there is at 
least this small area of sympatric distribution, then, and no evidence 
of intergradation in the available material, C. pygerythrus janetta and 
C. phayrei are here accepted as separate species. 

The form phayrei is separated from C. erythraeus atrodorsalis and 
C. e. zimmeensis by the apparently quite effective barrier of the Sal- 
ween River. Since phayrei also differs in color characters of the pel- 
age from both zimmeensis and atrodorsalis a great deal more than it 
does from C. e. sladeni or C. c. caniceps, it seems unquestionable that 
phayrei is of a distinct species, and no details for comparison with 
zimmeensis and atrodorsalis seem necessary here. 

C. phayrei has here been accepted as different at the species level 
from all tree squirrel forms to the south, west, east, and northeast 
of its known range that, like phayrei, ordinarily have only two pairs 
of functional mammae. It is separated from these other species to 
the south and west by the Salween River, and from a species to the 
northeast by the Irrawaddy River, and was formerly separated from 
two species to the east by a river barrier (the Irra-Sittang) . There 
remains only to ascertain the relationship of phayrei to the form im- 
mediately to the north and northwest. This is C. flavimanus shanicus 
which occurs in the area between the Salween and Irrawaddy rivers 
and has no barrier separating it from phayrei. 

The trenchant differences between phayrei and shanicus in pelage 
color are: (1) All four feet of shanicus are blackish agouti like its dor- 
sum, but those of phayrei are yellow or yellow-orange like its venter. 

(2) The ears of shanicus have the reddish rims, but those of phayrei 


are plain agouti like its back. (3) The ventral aspect of the tail of 
shanicus is as plain as the dorsal, but that of phayrei is ornamented 
by a longitudinal yellow midstripe. (4) The back of shanicus has a 
faintly expressed, broad, blackish, longitudinal band on the posterior 
two-thirds of the middorsum, but the back of phayrei is plain agouti 
like its sides, nape and crown. (5) The ventral pelage of shanicus 
is (somewhat obscurely) bisected by a longitudinal narrow stripe of 
agouti pelage, but that of phayrei has no such division. (6) the 
rostrum of shanicus shows more or less suffusion of rusty red, but 
that of phayrei is cool agouti like its crown. (7) The ventral pelage 
is generally a dull gray, but that of phayrei is a bright yellow-orange 
(sometimes with broad black margins extending between the fore 
and hind limbs). 

The first six of the above color characteristics of shanicus are gen- 
erally shared with C. f. gordoni, zimmeensis, atrodorsalis, thai, and 
other forms of C. fiavimanus to the east and south indicating a con- 
specific relationship with them while separating shanicus from phay- 
rei at the species level. Species distinction between these two is 
further indicated by both of them having been taken at precisely the 
same altitudes at three of the same localities, Maymayo, Gokteik, 
and Pyaunggaung, and there is a 100-mile north-south sympatry in 
their ranges indicated by mappable collecting localities. (See account 
of C /. shanicus for further details.) 

Thus, the form phayrei is clearly distinct at the species level from 
all other tree squirrels of its subgenus which surround it on every 
side, and if one accepts blanfordi and heinrichi as synonyms of it, 
phayrei is a monotypic species. 

Shortridge's field note on phayrei (in K. V. Ryley, 1914, p. 721) 
may shed light on the distributional relationship between phayrei 
and shanicus: "A particularly active species, its leaps from tree to 
tree almost rivaling those of Ratufa. Around Hsipaw town and 
Se'en, even more plentiful than . . . shanicus. Not observed at 
Maymyo." In contrast he found shanicus the most abundant squir- 
rel in Hsipaw State habitually near bungalows around Maymyo. 

What evidence we have suggests the possibility that C. f. shani- 
cus and C. phayrei may be in active competition with one another 
where their ranges meet or overlap and even that phayrei may be 
replacing shanicus in accordance with the "competitive exclusion 
principle" (Hardin, 1960). C ferrugineus also seems to be involved 
in this, but more locally and perhaps to a very minor extent making 
it three-way competition. 


Gallosciurus inornatus (Gray) 

Macroxus inornatus Gray, 1867, Ann. Mag. Nat. Hist. (ser. 3), 20, p. 282. 
Callosciurus imitator Thomas, 1925, Proc. Zool. Soc. London, 1925, p. 502. 

Types. — M. inornatus BM No., an old female from 
mountains in Laos, taken by C. Mouhote; imitator, BM No. 25.1.1.- 
68, adult male from Thai-Nien, latitude 22° N. on the Song Koi, 
Tonkin, 300 feet, collected February 26, 1924, by H. Stevens. 

Material examined, from Annam. — Hoi Xuan (CNHM), two; 
"Muong Sen," prov. Vinh (BM), one, (MNHN), one; Phu Qui 
(BM), three, (MNHN), eight; Lao Bao (MNHN), one; "Ipuing" 
(MNHN), one. 

Material examined, from Tonkin. — Phong Tho, 1000 feet (AMNH), 
one, (CNHM), one; Bac Tan Tray, 700 feet (AMNH), one; Lai 
Chau, 500 feet (AMNH), two, (ANSP), one, (CNHM), one, 
(USNM), three; M. Mouen (AMNH), two, (ANSP), eight; Na Hai 
(AMNH), two; Ba Nam Nung (CNHM), one; Muong Mo (CNHM), 
12; Muong Moun 1200 feet (CNHM), nine, (MCZ), one, (UMMZ), 
one; Muong Boum (CNHM), 11, (MCZ), two; Nong Lum (CNHM), 
one; Lieng San, 1500 meters (CNHM), six; Chapa (CNHM), four, 
(MCZ), two, (BM), 10; Pakha (CNHM), one, (BM), two. 

Material examined, from Laos. — Muong Yo, 2300 feet (CNHM), 
14, (USNM), two; Nap^, 2000 feet (BM), four; Pasa (AMNH), one; 
Xieng Khouang (BM), one; Don Qua (AMNH), one; Muong Koa 
[Khoua] (USNM), one; Nam Khueng (MCZ), two; Phu Kobo 
(MCZ), two; Lo Tiao (MCZ), two; Col de Taloun (MCZ), one; 
Phong Saly 4400 feet (MCZ), one, (UMMZ), one, (CNHM), three; 
"Tha Ngon," Vientiane (CNHM), one; Vientiane (USNM), one. 

Systematic history. — Gray's original description of inornatus fits 
the squirrel known since 1925 as imitator, quite well in every way 
excepting the last item on the tail: "Fur olive-grey . . . throat, inner 
side of limbs, and [venter] pale bluish grey, washed with whitish ; feet 
like back; tail longer than the body and head, colored like the back, 
with elongated white- tipped black hairs at the tip; hairs of the tail 
yellow, with three black bands ..." One of us (Tate) examined the 
type in 1951, noted it to be very much like the type of C. pygerythrus 
stevensi, but did not compare it with "imitator," and photographed 
and measured the skull. There is no doubt from the skull that this 
is Callosciurus, and there is no other Callosciurus species known from 
Laos which the type of inornatus reasonably could represent. Badly 
worn tails sometimes have the ends of the hairs broken off enough to 


reduce the count of black color bands, and tails with the hairs not yet 
grown out to full length do not show the full number of black bands. 

Thomas (1925) in describing imitator from Tonkin did not com- 
pare it with inornatus Gray and seemed unaware of the existence of 
inornatus, for he remarked the peculiarity of its possible relationship 
to the C. caniceps complex and noted its "superficial resemblance" to 
C. pygerythrus stevensi. Robinson and Kloss (1918) had overlooked 
inornatus in their nominal list of the squirrels of the Oriental Region. 

Pelage color. — Ventral pelage generally Light Violet Gray from 
chin to wrists and ankles. In the 68 specimens at Chicago Natural 
History Museum there is occasionally a collar of agouti intruded into 
this ventral pelage and a wedge of it extending posteriorly on the mid- 
line or a midstripe of it bisecting the bluish ventral pelage. The chin 
pelage is almost invariably bluish gray. Excepting for minor indi- 
vidual variations of lighter or darker, there seems to be no other vari- 
ation in the ventral pelage of this great series excepting the only 
summer specimen CNHM No. 32381 from Tha Ngon, Vientiane, 
Laos. This exception has a rather general suffusion of agouti through- 
out the ventral pelage. (Since the Vientiane specimen is from an 
area on the edge of the known species range and rather distant from 
sources of other material examined, one wonders whether the varia- 
tion might be geographic. A perhaps similar condition in an AMNH 
specimen from Na Hai, near Dien Bien Phu, suggests that the vari- 
ation may be individual.) 

The dorsal pelage is agouti, with two or often three light bands on 
the individual hairs, and Deep Olive (XL) in color. It varies aston- 
ishingly little in the large series at Chicago Natural History Museum. 
(There is some reddish suffusion the length of the dorsum, however, 
in 10/12 of the Muong Mo series and all 11 from Muong Boum, but 
it is poorly developed in the Lieng Sen six and Muong Mouen eight 
and virtually absent from the Phong Saly and Muong Yo series.) 
The ears are the same as the dorsal pelage; the feet are only a little 
more contrasty agouti. The tails are colored like the dorsal pelage 
and regularly but sparingly black- tipped (50 or 60 mm. black on 
light tipped hairs) throughout. The tail hairs regularly have five 
blackish bands on the individual hairs when the hairs are fully grown 
out, but one AMNH specimen has six. Table 10 provides measure- 
ments of two specimens which give some indication of the dimensions 
of this animal. 

This is a smaller squirrel than Callosciurus erythraeus hendeei with 
which it is almost completely sympatric (and which also shows re- 


markably little geographic variation in color characteristics) . Com- 
pare Figures 13 and 15. 

Diagnosis. — In describing imitator {=inornatus) Thomas noted its 
striking similarity to Callosciurus caniceps concolor far to the south in 
Malaya and to Callosciurus pygerythrus stevensi far to the northwest 
in Assam. C. inornatus has, however, a considerable number of 
somewhat varying pelage characteristics which together easily dis- 
tinguish it from C. p. stevensi: (1) The ordinary number of blackish 
bands on the tail hairs is five rather than four. (2) The tail is col- 
ored dorsally the same as the animal's back instead of paler and 
grayer. (3) The tail is lighter colored ventrally than dorsally but is 
yellowish brown instead of cool gray. (4) There is no flash mark of 
lighter color on the dorsal pelage of the hip as is common in stevensi. 
(5) The feet and forelegs are colored like the back instead of being 
notably more gray. (6) The subterminal light bands on the tail hairs 
are too short (ca. 2 mm.) and too yellow to give any special impres- 
sion as to the color of the tail hair tips, whereas in stevensi their white 
ness and length (ca. 3 mm.) give an impression of whitetipped hairs. 

Although C. caniceps concolor is most like C. inornatus in pelage 
characteristics, inornatus is geographically remote from concolor. On 
the other hand, C. inornatus is separated from the range of C. c. 
caniceps only by the Mekong River. Because of this proximity 
to caniceps it is important to consider the amount of difference be- 
tween inornatus and C. c. caniceps: (1) C. inornatus does not have a 
seasonal change of the dorsal pelage to orange but remains agouti 
gray (as do the subspecies of C. caniceps to the south of C. c. cani- 
ceps). (2) C. inornatus has agouti feet concolorous with the pelage 
of its dorsum instead of contrastingly pale gray. (3) C. inornatus 
has a vaguely delineated black pencil to its tail, obscured by whitish 
tips on the elongate hairs that compose it and varying amounts of 
light banding basal to the black on these hairs, unlike the crisply all 
black and sharply defined black tail tip of C. c. caniceps. (4) C. in- 
ornatus is substantially smaller than caniceps, greatest skull length 
of adults being about 51 mm. compared to 58 mm. in caniceps. 
(5) The rostrum and ear tips are colored like the back in inornatus 
(the tip of the rostrum tending a little toward black) instead of be- 
ing quite whitish as in C. c. caniceps. (6) C. inornatus seems gener- 
ally to lack the rather prominent longitudinal stripe of agouti which 
bisects the bluish gray venter of caniceps. 

Relationships to other species. — C. inornatus is almost wholly sym- 
patric with C. erythraevs hendeei, and was taken without any evidence 


of interbreeding in many of the same localities, and (when the ele- 
vation was given on the specimen tag) at the same elevation. 
Since inornatus is, furthermore, a smaller squirrel than hendeei, we do 
not question (nor has anyone) that they are specifically distinct. 
C. inornatus is separated from all of the subspecies of C. pygerythrus 
by two great barrier rivers, the Salween and the Mekong, and earlier 
the Irrawaddy must have constituted a third. It is shown above 
that inornatus generally has a good many differences in pelage color 
from the subspecies of C. pygerythrus that it most closely resembles. 

We recognize that the barrier strength of the great rivers may 
weaken to the north, that some other tree squirrel species are sep- 
arated into populations of no more than subspecific rank by any one 
of these river barriers in the north, and that C. pygerythrus stevensi is 
the northernmost subspecies of its species. These things do seem to 
support Ellerman and Morrison-Scott (1951, p. 488) in treating the 
species pygerythrus as the nearest relative of inornatus. On the 
amount of pelage color difference between inornatus and C. p. ste- 
vensi, the great gap in known distribution between them (a gap from 
which specimens of other tree squirrel species are nevertheless repre- 
sented in the collections available to us), and the interposition of the 
several major rivers as filter barriers, however, we cannot accept a 
subspecific relationship between inornatus and stevensi and conclude 
that these two forms are specifically distinct. 

Whether the species Callosciurus pygerythrus is indeed the closest 
relative of Callosciurus inornatus is to be questioned next. The dif- 
ferences between C. inornatus and C. c. caniceps described above are 
numerous and trenchant, but those between C. inornatus and C. cani- 
ceps concolor are exceedingly few. This close similarity between in- 
ornatus and concolor becomes significant in view of the following 
considerations which are emerging from the present study: (1) The 
Malay Peninsula below the Isthmus of Kra is a large mountainous 
area of little-differentiated tropical rainforest, virtually without sea- 
sons, and this region appears to provide many niches for tree squir- 
rels in which they remain conservative in pelage color. (2) Tonkin 
and upper Laos constitute an area which seems to have fewer niches 
for tree squirrels, but where they are similarly conservative in pelage 
color. (3) Thailand, excluding the peninsular portion, in strong con- 
trast with Malaya, contains large tropical lowland forest areas which 
differ sharply in annual rainfall, and have prominent alternating dry 
and rainy seasons, and in it tree squirrels develop the most fantasti- 
cally colored pelage. 


It seems reasonable to conceive that when CaUosciurus caniceps 
jfirst over-ran Thailand and spread throughout the area that is the 
present range of C. caniceps (Malaya to northern Thailand), it would 
then have had color characters much like concolor or inornatus 
throughout this range. Only two changes are needed then to achieve 
present conditions. (1) A stream piracy of the Mekong River by 
one of its own tributaries could have removed a substantial area 
from west to east of the Mekong River. A hypothetical example: 
the course of the Mekong might once have been through the valley 
which the Nam Tha now drains but was subsequently diverted to 
its present channel by head erosion of a recurved western tributary. 
This would have the effect of isolating a large population of this new 
species C. caniceps and freeing it to spread elsewhere east of the river. 
A population so separated from the species caniceps would reason- 
ably be expected to have spread to the limits now known for inor- 
natus and to have conserved rather well the pelage characters with 
which it arrived. (2) The population of species caniceps remaining 
west of the Mekong in the main part of Thailand would have evolved 
other pelage characteristics which would be the trenchant and extra- 
ordinary ones now observable in C. c. caniceps. 

That C inornatus thus may be directly derived from C. caniceps 
instead of C. pygerythrus does not appear to have been seriously con- 
sidered before, but may now be regarded as the more probable rela- 
tionship as a matter of minimum hypothesis. If inornatus is derived 
from species caniceps, there is an interesting niche reversal where 
sympatric with species flavimanus, for C. c. caniceps is the larger 
squirrel than C. /. tachin, and atrodorsalis and thai, but C. f. hendeei 
is the larger squirrel than imitator. 

CaUosciurus pygerythrus (Geoffroy St. Hillaire) 

Definition. — The species pygerythrus includes subspecies pyger- 
ythrus, janetta, owensi, mearsi, stevensi, hlythi, and lokroides, and 
their synonyms. These occur principally east of the Sittang and 
upper Irrawaddy valleys to the edges of the Indochinese Subregion 
as shown in Figure 16. 

Diagnosis. — The fully grown-out tail hairs of this species have 
but four blackish bands. There is a seasonal flash mark in the pel- 
age of the thigh which contrasts with the ordinary agouti dorsal 
pelage by being lighter (about cream color in most subspecies, but 
ochraceous buff in some) . These characters distinguish pygerythrus 
from sympatric species and adjacently allopatric ones. 




Fig. 16. Geographic distribution of the Irrawaddy squirrel, Callosciurus 
pygerythrus, as determined from material examined. Subspecies: A, pygerythrus; 
B, janetta; C, owensi: D, mearsi; E, stevensi; F, hlyihi; and G, lokroides. Dotted 
lines separate subspecies as known from specimens from the localities plotted but 
are not offered even as potential boundary lines between the subspecies. 

Relationships to other species. — This is a species of tree squirrel 
that is of smaller size than a species of tree squirrel generally attains 
in a hospitable geographic area which it occupies alone. Throughout 
virtually all of its range pygerythrus is sympatric with a larger spe- 
cies of tree squirrel, either Callosciurus erythraeus or C. ferrugineous. 
Compare their dimensions as indicated by Tables 11 and 4. This 
geographic range is constituted by those parts of Burma, India, and 
Pakistan which lie west of the Irrawaddy and Sittang rivers and east 
of the Brahmaputra River. An additional strip west of the Brahma- 
putra extends along the southern face of the Himalayas beyond the 


middle of Nepal. There is also a very small area that this species 
has invaded eastward across the Sittang Valley in the vicinity of 

It is shown in the account of Callosciurus phayrei that phayrei is 
the closest relative of Callosciurus pygerythrus and that through phay- 
rei, pygerythrus is next most closely related to Callosciurus caniceps. 
It is interesting that John Anderson (1879, p. 231) ventured much 
this same opinion about a definite interrelationship of pygerythrus, 
phayrei, and caniceps, from examination of a substantial series, but 
he was speculating that they are all one species. 

Intraspecific variation. — In subspecies lokroides the term "winter 
pelage" may be perfectly acceptable to distinguish the duller pelage 
of winter from the very much brighter pelage of summer. However, 
Raven collected subspecies mearsi in dull "winter" pelage without 
flash marks on the hip in March at several localities directly across 
the Chindwin River from where he was collecting subspecies owensi 
at the same time in bright "summer" pelage with flash marks. Short- 
ridge (in Wroughton, 1916, p. 293) remarked no pronounced differ- 
ence in the character of the forest on the two sides of the Chindwin 
River at these places, between Hkampti and Tamanthe. The pelage 
of two subspecies being out of phase across the river barrier raises 
some question as to whether the terms "winter" and "summer" pel- 
age are in fact suitable terms to be applied to these subspecies of lower 
latitude and elevation. Perhaps "eclipse" pelage would be better ter- 
minology for the dull pelage, where a distinctive alternation occurs. 

We have examined and compared summer and winter specimens 
of the subspecies janetta, which occupies the dry scrub forest of the 
dry zone of central Burma, but the differences between these are so 
slight that one can hardly claim there are seasonally bright and dull 
pelages in janetta at all. The flash mark on the hip is present in all 
specimens of janetta. Part of what we recognize here as mearsi west 
of the lower Chindwin and lower Irrawaddy also occurs in the ex- 
treme dry zone of central Burma. Material of mearsi collected in 
winter in this zone is, of course, distinct from janetta, but surprisingly 
resembles it in having a flash mark on the hip; whereas the large 
winter season series of mearsi from the deciduous forest zone seems 
to be in dull seasonal pelage and has no flash mark on the hip. Thus, 
we have a species which as a whole seems to alternate dull pelage 
(and no flash mark) with bright pelage (and flash mark) . But within 
this species is one subspecies, janetta, and part of another, mearsi, 
that unlike the rest of the species occur in the scrub forest of a nearly 

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desert area. Both of these have flash marks in the winter pelage and 
other reductions in the amount of seasonal difference in color of pelage. 
The pelage characters which distinguish the subspecies of the spe- 
cies pygerythrus are relatively inconspicuous ones, and in order to 
distinguish them we were obliged to write down lists of the color 
characters of each series and test them empirically to see what does 
distinguish each alleged subspecies from the others geographically 
adjacent to it. This procedure resulted in valuable, recorded de- 
scriptions and diagnoses easily included in the present paper. The 
characters of subspecies of other species have generally been better 
known and easier to observe and remember, and such time-consuming 
procedure as employed with pygerythrus was not necessary nor prac- 
tical for so extensive a study as the present one. 

Callosciurus pygerythrus pygerythrus (Geoffroy St. Hilaire) 

Sciurus pygerythrus Geoffroy St. Hilaire, 1832, Mag. Zool., Paris, fifth unnum- 
bered page for Pis. 4, 5, and 6, CI. 1; see also 1834, Geoffroy St. Hilaire, in 
Belanger, Charles, Voyage aux Indes-Orientales. Zool. p. 145 in text, 
pL 7 in atlas. 

Ti/pe.— MNHN No. 1829-286(294), adult from forest of Syriam, 
near Pegu, Burma, collected by Charles Belanger. 

Material examined, all from Burma. — "Camp Pinmezali," 850 
feet, Pegu Yoma (AMNH), one; "Yetho River," 100 feet, Pegu Yo- 
ma (AMNH), five; "Kathipinzan," 100 feet, Pegu Yoma (AMNH), 
two; Hmawbi, 40 km. north of Rangoon (AMNH), one; Shandaw, 
125 km. north of Rangoon (AMNH), one, 30 km. north of Prome 
(AMNH), six; Zaungtu, 30 miles north of Pegu (CNHM), one; 
Tamabin, 200 feet, 25 miles north of Pegu (CNHM), one, (BM), 
three; South Zamaya Reserve, 45 miles north of Pegu (BM), one; 
20 miles north of Toungoo (BM), one; 30 miles north of Toungoo 
(BM), one; 40 miles north of Toungoo (BM), two; Rangoon (BM), 
four; "Tenasserim" (BM), one; Pegu (BM), three. 

Pelage color. — Description from series in American Museum of 
Natural History: (1) The dorsal pelage is about Olive Brown (XL) 
from rostrum almost to tip of tail, and hardly lightens on the ears 
or sides. (2) Dorsal pelage of the feet is agouti but is appreciable 
lighter than dorsum. (3) The sides of the neck are like the feet. 
(4) There is a short black tip to the tail. (5) Blackish bands on indi- 
vidual tail hairs are as many as four. (6) Ventral pelage is Cinnamon 
Rufous (XIV) but with the chin, throat, and breast Pale Ochraceous 
Buff to Light Ochraceous Buff (XV). (7) Perineal pelage does not 
differ, or a small amount of it is lighter. (8) The midventral, longi- 


tudinal, tail stripe is pale and feebly expressed in most (but brilliant 
and strongly expressed for full length of tail in the Shandaw specimen) , 

Diagnosis. — The subspecies C. p. pygerythrus is distinguished 
from the known adjacent conspecific subspecies by the above char- 
acteristics as follows: From C. p. janetta by Nos. 1, 2, and 6. 

Discussion. — Our measurements in Table 11 and photographs 
studied of the skull of the type specimen leave us in no doubt that 
its nearest relative is janetta. The type has no flash mark on the hip 
and the two specimens in the Chicago Natural History Museum col- 
lection, which were both taken in January, have none. Since the 
American Museum of Natural History material, which was taken 
April 28, May 7-11, and September 10-16, all has the flash mark 
present, it becomes evident that this purely tropical subspecies has 
a seasonal change of pelage resembling those of the more northern 
subspecies in that the flash mark is present on the hip during the 
summer (rainy) season and absent during the dry winter. In this 
C. p. pygerythrus differs from C. p. janetta, for janetta has the flash 
in both summer and winter. 

Interestingly enough the American Museum of Natural History 
material of C. p. pygerythrus shows striking evidence of molt in April 
and May as well as September, suggesting two molts a year as the 
mechanism by which the observed difference in seasonal pelage is 
brought about. 

The material from 30 kilometers north of Prome is distinctly 
intermediate in pelage color characters between C. p. pygerythrus 
and C. p. janetta. Prome is about where de Terra (1944, fig. 4) draws 
the line between the vegetation zone "Delta and Swamp Forest" 
on the south and "Dry Scrub Forest" on the north. Since all of the 
locality records for C. p. janetta are to the north in the Dry Scrub 
Forest Zone and all the C. p. pygerythrus localities found are in the 
zones of Delta and Swamp Forest, Deciduous Forest, and Rain- 
forest which lie south of the Dry Scrub Forest Zone, the hypothetical 
line along which C. p. pygerythrus and C. p. janetta are presumed to 
intergrade approximates the southern margin of the Dry Scrub 

Callosciurus pygerythrus janetta (Thomas) 

Sduriis pygerythrus janetta Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, 
p. 203. 

Type.— BM No., adult male from Mandalay, 200 feet, 
collected June 28, 1913, by Guy C. Shortridge. 


Material Examined, all from Burma, and all from east of the 
Chindwin River. — Okma (AMNH), seven; Mingun, west of Sagaing 
(CNHM), two, (BM), eight; "Gada," Shwebo (BM), one; "Kyon- 
daw," Kawlin, Shwebo (BM), one; "Kayikthin," Shwebo (BM), one; 
Mandalay (BM), six; "Myitnge River," 500 feet, Maymyo (BM), one; 
Monywa (BM), one; "Nyoring Vintha," east bank of Irrawaddy 
(BM), one; "Pyawbwe," 700 feet (BM), two; Wuntho, Shwebo (BM), 
one; Yin, Lower Chindwin River (BM), seven; Mt. Popa, 600 meters 
(AMNH), three, (BM), five; 20 km. north of Meiktila (AMNH), one. 

Pelage color. — (1) From Okma, Mt. Popa, and Meiktila material, 
janetta is seen to possess agouti dorsal pelage about Grayish Olive 
(XLVI) in general aspect. (2) This color is exceedingly uniform 
from back to crown, ears, tail, sides, or legs. (3) There is a rather 
small Cream Color flash mark on the hip in the dorsal pelage. (4) 
The edge of the flash mark is contiguous with the edge of the ventral 
pelage, and not separated from it. (5) All four feet are cream colored 
on the digits and often on the whole foot. (6) There is a rather 
inconspicuous short black tip to the tail. (7) There are up to four 
blackish bands on the tail hairs. (8) The ventral pelage is about 
Light Ochraceous Buff (but in extreme individuals is Pale Ochraceous 
Buff or Ochraceous Buff), which is slightly huffier than the flash 
mark. (9) The ventral pelage is gray on the basal portion of the 
individual hairs in a broad band across the abdomen. (10) The 
mid-ventral pelage of the tail is about Ochraceous Buff proximally for 
about the middle third of its width, but diminishing distally both in 
width and intensity of color. 

Diagnosis. — Callosciurus pygerythrus janetta may be distinguished 
from geographically nearest conspecific subspecies by the above enu- 
merated color characters as follows: from C. p. owensi by nos. 1, 3, 
4, 5, 8, and 10; from C. p. mearsi (as known in ecHpse pelage) by 
nos. 2, 5, and 6; from C. p. pygerythrus by 1 and 8. 

Discussion. — The difference between janetta and C. phayrei with 
which it is apparently sympatric in at least a small area about Man- 
dalay and Ava, are detailed in the account of the species phayrei. 
Since at Mandalay janetta comes fairly close to the range of C. fiavi- 
manus shanicus, the differences between these two forms and possibil- 
ity of intergradation need to be considered. C. f. shanicus differs from 
C. p. janetta in pelage color characters enumerated above for the latter 
as follows: Nos. 1, 2, 3, 5, 7, 8, 9, and 10. Pelage characteristics that 
shanicus possesses in common with C. jiavimanus gordoni and /or 
C. jiavimanus atrodorsalis and does not share with janetta are: (1) 


Margins of ears are reddish. (2) Rostrum is reddish. (3) There is 
a faint but distinct indication of a broad, blackish band on the pos- 
terior two- thirds of the mid-dorsum. (4) The dorsal pelage of the 
feet is blackish agouti. (5) Although barely distinguishable there is 
an indication of a band of agouti longitudinally bisecting the ven- 
tral pelage. 

There is also a size difference of significance between shanicus and 
janetta which may be seen, for example, in the hind foot measure- 
ments of the samples of these made by the same collector, Gerd 
Heinrich: shanicus, 45, 45, 47, 47 mm.; janetta, 40, 40, 40, 43 mm. 
In all, the differences between janetta and shanicus are thus obviously 
greater than subspecific. 

We are unenthusiastic about accepting janetta as a single sub- 
species. Final decision has had to be made with only the American 
Museum of Natural History material at hand, and although slight 
but consistent differences are apparent in this between the samples 
from the populations separated by the Irrawaddy River, the sample 
from north of the river is all from one locality. When some future 
student is able to investigate this with material better representing 
the range of janetta on both sides of the river, he may find it necessary 
to recognize them as two subspecies. 

Callosciurus pygerythrus owensi (Thomas and Wroughton) 

Tomeutes similis owensi Thomas and Wroughton, 1916, Jour. Bombay Nat. 
Hist. Soc, 24, p. 236. 

Type.— BM No., old female from Minsin, 450 feet, 
upper Chindwin River, Burma, collected August 15, 1914, by G. C. 
Shortridge and S. A. Macmillan. 

Material examined, all from Northern Burma, between the Chind- 
win and Irrawaddy rivers. — Lonkin (AMNH), one; Mansum 
(AMNH), two; Pumsin (AMNH), one; Limpa (AMNH), two; 
Kaunghein (AMNH), three; Moklok (AMNH), four; Phawzaw 
(AMNH), two; Tamanthe (AMNH), two; Hkamti (BM), one; 
Kauktaung (BM), one; "Northern Burma" (AMNH), one. 

Pelage color. — Seventeen American Museum of Natural History 
specimens come from eight localities, two of the eight straddling the 
type locality and about five miles from it. These were taken between 
January 14th and March 18th. (1) The dorsal pelage is agouti and in 
the mid-dorsal, more reddish portion appears about Cinnamon Brown 
(XV). (2) The sides and hind legs are agouti but not reddish and 


appear about Light Brownish Olive (XXX). (3) The reddish color 
of the back diminishes very gradually anteriorly and is generally 
faint on the nape and absent from the crown and ears which are 
about the color of the sides but slightly darker. (4) There is a buffy 
eye ring. (5) The pelage on the foot is the same color as that on 
the leg. (6) The reddish color of the middorsum continues pos- 
teriorly for most of the length of the tail. (7) Buffy sub terminal 
bands on the tail hairs provide a noticeable buffy fringe. (8) The 
tip of the tail is black for about the distal half of the terminal hairs. 
(9) The flash mark on the hip is Ochraceous-Buff (XV) . (10) The 
flash mark (in all but one case) is separated from the ventral pelage 
by some agouti dorsal pelage. (11) The ventral pelage is longitudi- 
nally divided by a thin midstripe of dark pelage. (12) The ventral 
pelage has light color, or long light tips which give it an overlying 
color, of Light Ochraceous-Buff (XV), but dark gray bases to the 
hairs, everywhere but the axils and groin and sometimes the forelegs. 
(13) The perineal pelage of adult males is more richly buffy colored 
than the general ventral pelage of the same animal. (14) The ven- 
tral color of the tail lacks the reddish dorsal infusion and has no 
proximal midventral intrusion of strong buffy color. 

Diagnosis. — Callosciurus pygerythrus owensi is distinguished by 
the above described winter pelage characteristics from C. p. stevensi 
to the north by nos. 5, 11, and 12; from C. p. mearsi to the west by 
nos. 1, 6, 7, 8, 9, and 13; from C p. janetta to the south by nos. 1, 2, 
5, 6, 7, 9, 10, 11, 12, and 13. 

Discussion. — The winter pelage of subspecies owensi suggests 
strong affinities between owensi and stevensi in the dark-colored dor- 
sum with a reddish middorsal infusion, in the presence of a flash 
mark on the hip in winter pelage, in the rich color of the flash mark, 
in the presence of gray bases to the ventral pelage of the hind legs, 
breast, and throat. Some direct interbreeding with mearsi despite 
the Chindwin barrier is certainly suggested by the divided venters 
seen in the northern mearsi material. 

The type description by Thomas and Wroughton is of an animal 
in summer pelage, and since we cannot provide better detail, we 
quote it. "General ground color above essentially the same dark 
grizzled olive gray as in [summer pelage of lokroides], but the whole 
back strongly suffused with deep rufous, becoming stronger poste- 
riorly, the loins practically grizzled 'chestnut,' passing without con- 
trast into the ferruginous of the hip-patch. Head, neck, shoulders 
gray. Under-surface buffy, a more or less distinct grizzled gray line 


running down the center from throat to belly; a small central patch 
in inguinal region whitish. Inner sides of forearms whitish, of legs 
strong buffy. Hands and feet grizzled gray with or without buffy 
intermixture. Tail coarsely grizzled gray with but slight buffy suf- 
fusion; the terminal hairs black." 

Callosciurus pygerythrus mearsi (Bonhote) 

Sciurus lokroides mearsi Bonhote, 1906, Ann. Mag. Nat. Hist. (ser. 7), 18, 

p. 338. 
Tomeutes mearsi bellona Thomas and Wroughton, 1916, Jour. Bombay Nat. 

Hist. Soc, 24, p. 420. 
Tomeutes mearsi virgo Thomas and Wroughton, 1916, Jour. Bombay Nat. 

Hist. Soc, 24, p. 421. 

Types. — S. I. mearsi, BM No., old female from Chinbyit 
on a west bank tributary of the Chindwin River, collected January 16, 
1906, by A. Mears; bellona, BM No., old male from Kin, 
Lower Chindwin River, Burma, collected June 18, 1914, by G. C. 
Shortridge; virgo, BM No., adult male from Tatkon, near 
Kindat, Upper Chindwin River, Burma, collected July 5, 1914, by 
G. C. Shortridge. 

Material examined, from Burma, and all from west of the Chind- 
win River.— Mt. Victoria, Pakkoku Chin Hills (AMNH), one; 
Dudaw-Taung, Pakkoku Chin Hills (AMNH), one; Kyundaw, 30 
km. northwest of Seikpyu (AMNH), six; Kalewa (AMNH), 11; 
Mawlaik (AMNH), three; Kindat (AMNH), two, (BM), four; 
Homalin (AMNH), 20, (BM), four; Tempao (AMNH), eight; Ta- 
manthi (AMNH), one, (BM), four; Phawzaw (AMNH), one; Kin 
(BM), 11; Okma (AMNH), three; Moklok (AMNH), one; "Ma- 
gaung," lower Chindwin (BM), one; Tatkon (BM), seven, (CNHM), 
two; "Aingma," lower Chindwin (BM), one; Sangau, Lushai Hills, 
(CNHM), four; Kabaw Valley, 20 miles west of Kindat (BM), two; 
Chaung, 20 miles n.n.w. of Kindat (BM), one. 

Material examined, from Assam. — Kohima, Naga Hills (CNHM), 
two; Karong, Manipur (CNHM), one; "Therighat," Manipur (BM), 
one; Mokokchung, 4500 feet, Naga Hills (BM), one; Margherita 
(BM), two; "Sangrachu," 3000 feet, Naga Hills (BM), one; Gola- 
ghat, 400 feet (BM), five; Thotal [Thobal], Manipur (BM), one; 
"Naga Hills" (BM), one; "Namdang" Naga Hills (BM), one. 

Pelage color. — The following description is from the American 
Museum series of 55 specimens from 11 localities between Moklok 
and Mt. Victoria 325 miles to the south. The ones from Chindwin 


River localities are all in eclipse pelage having been taken between 
March 16 and April first. The ones from farther south at Kyundaw 
and Dudaw-Taung were collected between January 28 and 29, but 
the Mt. Victoria specimen was taken on July 2nd and is in summer 
pelage. (1) The agouti dorsal pelage is generally about Buffy Brown 
(XXX) but on some few specimens is Light Brownish Olive. (2) In 
the winter pelage there is a pronounced tinge of Isaballa Color in the 
agouti dorsal pelage, most intense across the shoulders (and quite 
prominently so when series are laid out) but often coloring the sides, 
also (quite like character no. 2 of lokroides). (3) There is an eye ring 
buffy enough in color to contrast well with the agouti pelage of the 
face and often wide enough to be quite conspicuous. (4) The flash 
mark on the hip is white. (5) The ventral pelage is light-colored 
near the tip but, when fully grown out, is contrastingly gray near the 
bases of the hairs at least on the area covering the abdomen. (6) 
There is a pair of contrastingly buffy patches of pelage on the 
groin. (7) Perineal pelage, if it is distinct from the other ventral 
pelage, is only whiter. (8) The number of blackish bands on fully 
grown out tail hairs is four. (9) In winter pelage the proximal, mid- 
ventral pelage of the tail is quite buffy and this color may extend 
midventrally out the tail for much of its length or not. (10) There 
tends generally to be a rather obscure dash of reddish color in the 
tail near its tip. 

Discussion. — There is much undescribed geographic variation in 
mearsi as it is accepted here. If Thomas and Wroughton (1916, 
pp. 419^22) had presented adequate descriptive and other data on 
the collections on which they reported, it might have been possible 
to understand the variation in summer pelage and to present a co- 
herent systematic account of it. One of us (Tate) has subsequently 
examined that summer pelage material in the British Museum and 
was apparently quite unimpressed by the alleged differences between 
mearsi, virgo, and bellona. Neither of us finds any characteristics in 
the eclipse pelage to support recognition of these named forms. How- 
ever, material in eclipse pelage from the west bank of the Chindwin 
at Tempao, Phawzaw, and Moklok is characterized by darker ven- 
tral pelage which is longitudinally bisected by a thin agouti stripe. 
This material represents the three northernmost Chindwin River 
localities of mearsi. The southernmost localities are represented by 
midwinter material from Kyundaw and Dudaw-Taung and a sum- 
mer specimen from Mt. Victoria. All of the Kyaundaw six are pro- 
vided with a flash mark on the hip, and are very whitish beneath. 


The summer specimen from nearby (30 to 40 miles) Mt. Victoria has 
notably browner dorsal pelage and a larger flash mark. These dif- 
ferences suggest that at Kyundaw there are flash marks in the winter 
pelage or that in this lower latitude there may be but one instead of 
two annual molts. 

Diagnosis. — Callosciurus pygerythrus mearsi differs from C p. 
lokroides by the above described characters in nos. 3, 4, and 7; from 
C. p. stevensi by nos. 1, 2, 4, 6, 9, and 10; from C. p. blythi by nos. 1, 

2, and 7; from C. p. owensi by nos. 1, 2, 4, 6, 7, 9, and 10; from C. p. 
janetta by nos, 2, 3, 6, and 10; and from C. p. pygerythrus by nos. 2, 

3, 6, 7, and 10. 

Habits. — Morris (1936) records the following observations on 
mearsi from the field notes of Charles McCann: On p. 670 at Kalewa 
where the expedition took 11 mearsi, they "... were apparently 
feeding on the fruit of Calycopteris floribunda." On p. 668 across 
the river from Homalin where 14 specimens of mearsi were taken 
one day, "The scrub jungle in parts seemed to be alive with this 

Callosciurus pygerythrus stevensi (Thomas) 

Sciurus stevensi Thomas, 1908, Jour. Bombay Nat. Hist. Soc, 18, p. 246. 

Type.— BM No., adult male, from near Beni Chang, 
Abor-Miri Hills, 4000 feet, Assam, India, collected February 19, 1906, 
by H. Stevens. 

Material examined, from Burma, west of the Chindwin River. — 
Limpa (AMNH), one; Hahti (AMNH), two; Haibum (AMNH), six; 
Dalu (AMNH), one; 20 miles north of Hkamti on Chindwin River 
(BM), one. 

Material examined, from Assam, India. — Benichang, 4000 feet, 
Miri and Abor Hills (BM), one; Kalek, Abor Hills (BM), one; Sadiya 
(BM), two; Ledo, 200 feet (BM), one; 22 miles east of Ledo (USNM), 

Pelage color. — The subspecies stevensi is quite distinct and appar- 
ently fairly stable in the diagnostic color. (1) The agouti dorsal 
pelage ranges from Light Brownish Olive (XXX) to Cinnamon 
Brown (XV). (2) The ventral pelage is undivided. (3) The ven- 
tral pelage is composed of hairs with whitish tips that overlie but 
fail to hide the dark-gray hair bases, and the general effect is a bluish 
gray, about Pale Quaker Drab (LI) . (4) A hip mark is clearly pres- 
ent in four of the American Museum of Natural History series (10) 


and is Apricot Buflf (XIV). (5) The head, hind legs, and ears are 
colored like the back, which pales slightly on the sides. (6) The 
front legs and all the feet are notably grayer than the dorsum. In 
a few specimens the difference is only slight but in series it is still 
fairly obvious. (7) The number of blackish bands on the fully grown 
out tail hairs is four. (8) Dorsally the tail is colored paler and grayer 
than the pelage of the back. (9) The tail is still a cooler gray below 
than it is above because the light bands on the ventral tail hairs are 
whitish instead of pale buff. (10) The subterminal light bands on the 
tail hairs are light enough and long enough (ca. 3 mm.) to give the 
gross impression that the hairs are white- tipped. 

Diagnosis. — The characteristics of stevensi described and num- 
bered above distinguish stevensi from the other subspecies of pyge- 
rythrus that are geographically adjacent to it as follows: C p. stevensi 
is distinguished from owensi to the east and south across the Chind- 
win River by nos. 2, 3, 8, 9, and 10; from mearsi to the south by 
nos. 1, 2, 3, 4, 6, 7, 8, and 9; from hlythi to the southwest by nos. 3, 
4, 5, and 9; and from lokroides to the west by nos. 1, 2, 3, 6, 8, 9, 
and 10. 

DisciLSsion. — It needs to be pointed out that the Dalu specimen 
is the only one of the (AMNH) series taken on the east side of the 
Chindwin River and that the Dalu specimen is an intergrade with 
owensi, for it possesses incipient tawny groin patches, an even brighter 
patch (about Warm Buff) of perineal hair, faintly lighter and very 
faintly buffy ventral pelage on the body, and its tail does not con- 
form to characters 8, 9, or 10 above. 

The specimen of stevensi from Limpa is from the west bank of the 
river, and this locality is the most southern for the subspecies. Al- 
though the ventral pelage is typical in the Limpa specimen, there is 
a hint of intergradation with mearsi in the obvious failure of the tail 
to conform with stevensi characteristic number nine above. 

The similarity between Callosciurus pygerythrus stevensi and Cal- 
losciurus inornatus and their differences are described in the account 
of that species. 

Robinson (1913, p. 95) reports a series of stevensi from the Abor 
Hills: three from Balek, about 4 miles west of Pasighat on the west 
side of the Dihang; four from Rotung just south of the Dihang at 
95° 10' east longitude; one from near Kalek, which is about three 
miles south of Rotung, and one from five or ten miles south of Kalek. 
Robinson (p. 89) describes this series as "very uniform and [it] agrees 


well with the description of the type ..." The present distribution 
of Callosciurus pygerythrus stevensi across the great river barrier of 
the Brahmaputra from the Naga Hills to the Abor Hills west of the 
Dihang appears to constitute a piece of zoological evidence support- 
ing the hypothesis that the Dihang was formerly a tributary of the 
Chindwin and has been captured by the Brahmaputra (see Chhibber, 
1934, p. 33). 

Callosciurus pygerythrus blythi (Tytler) 

Sciurus blythi Tytler, 1854, Ann, Mag. Nat. Hist. (ser. 2), 14, p. 172. 

Cotypes.—BM No., adult males, both from 
Dacca, East Pakistan, collected by R. C. Tytler. 

Material examined, all from Assam, India, and East Pakistan.^ — 
Umran, Khasia Hills (AMNH), two, (CNHM), one; Nongpoh, 
Khasia Hills (AMNH), one; Burnihat, Khasia Hills (AMNH), 
seven, (CNHM), one; Palasbari (UMMZ), two, (CNHM), four; 
Bamanigoan (UMMZ), one; Cherrapunji, Khasia Hills (CNHM), 
one; Mawlyngkueng, Khasia Hills (CNHM), two; "Dilkoosha," 
Cachar (BM), one; "HatikhaH," 1500 feet, Cachar (BM), one; 
"Lanka," 400 feet, north Cachar (BM), two; "Laitkynsao," 2500 
feet, Khasia Hills (BM), one; "Matunga River," 300 feet (BM), 
three; Tura, 1400 feet, Garo Hills (BM), one; Duragiri, 3000 feet, 
Garo Hills (BM), one; "Rangapani," 400 feet, Garo Hills (BM), 
one; Angarakhata, 300 feet, N. Kamrup (BM), two; Dacca, E. Paki- 
stan (USNM), one; 20 miles north of Dacca, E. Pakistan (USNM), 

Pelage color. — The one topotype and one near-topotype (USNM), 
and the material in the American Museum of Natural History are 
described as follows: (1) The mid-dorsal pelage is grossly about 
Olive-Brown (XL) grading into Buffy Brown on the sides. (2) The 
legs and feet are grayer and cooler than the back and sides. (3) 
The head is darker, grayer, and of a more contrasting agouti than 
the back and sides. (4) The whitish venter is about Pale Ochraceous 
Buff except for a substantial spot in each axil as well as in each groin, 
which are the richer color Light Ochraceous Buff. (5) There are as 
many as four blackish bands on the tail hairs, and the elongate hairs 
on the tip of the tail do not contrast notably in color with the rest 
of the tail. (6) The hip mark is present and prominent on alm^ost 
every specimen and is whitish or Cream Color. (7) The under side 
of the tail is Light Ochraceous Buff to Apricot Buff in the middle 


one-third or one-fourth of the tail's width, and in some adult speci- 
mens this color extends full strength for the whole length of the tail. 

Diagnosis. — C. p. blythi is distinguished from its geographically 
nearest conspecific relatives by the above-numbered color charac- 
teristics as follows: lokroides by 3, 4, and 6; stevensi by 4, 6, and 7; 
and mearsi by 3 and 4. 

Discussion. — The USNM material recently obtained from the 
vicinity of Dacca appears to confirm this locality, which is about 
65 miles out on the Ganges-Brahmaputra delta west of any hills, as 
the type locality of the subspecies blythi. Khajuria {in Moore, 1960, 
p. 8) remarks that he has taken this form at other localities in the 
delta near the Garo, Khasia, and Jaintia hills in Assam. It is, thus, 
apparently the only diurnal squirrel of the Indochinese Subregion 
which has come down off the jungle-clad hills of Assam into the plains 
of the delta as if capable of crossing the Garo-Rajmahal Gap into 
the Indian Subregion. Tytler (1854) in reporting on the fauna ob- 
served while marching from Barrackpore (20 miles north of Cal- 
cutta) via Jessore and Faridpur to Dacca, remarked that during the 
20 days of this march Sciurus palmarum [Furnamhulus pennanti] be- 
came scarce and was not seen at all east of the Puddur River. He 
said it was unknown at Dacca but "it is replaced by a fine species 
found in the jungles and by no means uncommon. . . . They are by 
no means timid, but after being once disturbed and alarmed, run and 
hide among the branches. ..." The details of distribution of blythi 
on the delta in relation to that of Funambulus pennanti would be 
quite interesting to know. 

Callosciurus pygerythrus lokroides (Hodgson) 

Sciurus lokroides Hodgson, 1836, Jour. Asiatic Soc. Bengal, 5, p. 232. 
Sciurus assamensis Gray, 1843, List Spec. Mammal. Brit. Mus., p. 143, 
Macroxus similis Gray, 1867, Ann. Mag. Nat. Hist. (ser. 3), 20, p. 281. 

Types.— S. lokroides, BM No., old female. No. 43.1.12.- 
59, young male, both from Nepal; BM No., adult male, 
from Nepal. Probable cotypes: Leiden specimen a (no. 13364) Hodg- 
son no. 28, old individual from Nepal; Leiden specimen b (no. 13365), 
Hodgson no. 62; assamensis, BM No., adult, from As- 
sam; similis, BM No., and BM No., adult male, 
both from Nepal. 

Material examined, from Nepal. — Sisagarhi, 5875 feet (AMNH), 
two; "Nepal" (BM), five; Naggenjung, 6000 feet, Katmandu 
(AMNH), six; Amlekhganj (AMNH), one; "Lokoripawa," 8000 feet 


(BM), two; Hetora (AMNH), one; Aromari (BM), one; Sunuchuru 
(BM), one; Riri Bazaar, W. Nepal (CNHM), one; Hetwada (BM), 
one; Katmandu, 4500 feet (BM), four; Nawacot, 7000 feet (BM). 

Material examined, from Sikkim. — Lingtam (UMMZ), one, 
(CNHM), 12; Dikchu, 2000 feet (CNHM), one, (BM), one; Toong 
(CNHM), one; Rongli, 2700 feet (BM), four; "Sikkim" (BM), two; 
Rao Dala, 4500 feet (BM), two. 

Material examined, from Bengal. — Sangsir (CNHM), four; Gop- 
aldhara, 5000 feet, Rungbong Valley (BM), two; Mangpu, 3500 feet 
(CNHM), nine; Sevoke, 500 feet (BM), five; Darjeeling (BM), one; 
Pashok, 3500 feet, Darjeeling (BM), three; Mongtu, hills, 3850 feet, 
west of Tista River, Darjeeling (BM), four; Nanh laing, R. S. (BM), 
one; Machi, Manipur (BM), one; "Pachoe," 3500 feet, Darjeeling 
(BM), two; Haraincha, Morang, East Terai (BM), one. 

Material examined, from Bhutan. — Bharnabhare, 600 feet, Bhu- 
tan Duars (BM), two; Bhotan (BM), four; Hasimara, 600 feet (BM), 

Pelage color. — Callosciurus pygerythrus lokroides taken March (1), 
April (8), and May (1) may be described from the American Museum 
material as follows: (1) It has agouti dorsal pelage which is about 
Drab or Hair Brown (XLVI) which seems to have but two light 
bands on each guard hair. (2) The sides appear to be bufRer as a 
result of diminishing in the total amount of black on the parts of 
hairs showing at the surface, and have the look of C. c. caniceps in 
the gray seasonal pelage. (3) The pelage of the head, ears, legs, and 
dorsum of tail does not differ appreciably in color from that of the 
back. (4) All four feet are appreciably cooler and grayer than the 
middorsum. (5) The flash mark on the hip (present only in the one 
summer pelage specimen) is in the area of the dorsal pelage of the 
leg, but borders on the ventral pelage. (6) The flash mark of the hip 
is Ochraceous-Orange. (7) The hairs of the tail have four blackish 
bands each. (8) The tail is concolorous to the end in the dorsal view 
(i.e., has no black or other contrasting color at the tip) . (9) The ven- 
tral pelage is composed of hairs with long light tips but dark gray 
bases (at least on the body), and there is a tendency for the light tips 
to be buffy (abdominally) instead of whitish. (10) There are a pair 
of buffy patches of pelage on the groins (which are indistinguishable 
in individuals that have especially buffy general color of the venter) . 
(11) Buffy perineal pelage is present in adults, the same color as the 
groin patches and in males often continuous with them. (12) Prox- 
imally the mid-ventral pelage of the tail is buffy. 


Diagnosis. — The characteristics described and numbered above 
distinguish lokroides from geographically adjacent other subspecies 
of pygerythrus as follows: from stevensi by nos. 1, 2, 3, 5, 8, 9, 10, 11, 
and 12; from blythi by nos. 3 and 6; from mearsi by nos. 6 and 11. 

Habits. — A collector reports field observations as follows: "Found 
at low elevation. At Rongli it was very partial to oranges, doing 
much damage to the crop. The 'dray' is a collection of grass and 
sticks, placed high up in a tree. This squirrel is found in heavy for- 
est, and near villages and may often be seen on the ground searching 
for food. Considering the size of the animal its call is sometimes very 
loud." (C. A. Crump in Wroughton, 1916a, p. 487.) 



Type species. — Tamiops maritimus hainanus J. A. Allen. 

Definition. — The genus Tamiops is constituted by the species 
mcclellandi, rodolphei, swinhoei, and maritimus, and occurs virtually 
throughout the Indochinese Subregion as suggested by the maps of 
its distribution in Figures 18, 19, and 20. 

The little tree squirrels of this genus range from Malaya to Hopeh, 
China, about the same range as that of Dremomys and Sciurotamias 
combined. This is a range of almost 40° of latitude and about 3000 
miles. This genus occurs at low elevations in tropical rain forest in 
some parts of its range and up to 12,000 feet elevation in the moun- 
tains, and it is reputed to be a quiet, secretive squirrel. It has a 
prominent basic dorsal pattern of five longitudinal black stripes sep- 
arated by four lighter stripes, and these are subject to some geo- 
graphic and seasonal variations in number and intensity. The face 
has a prominent light stripe passing from the dorsal surface of the 
rostrum posteriorly beneath the eye. The tail as well as the stripes 
is somewhat chipmunk-like, being quite short-haired for a tree squir- 
rel. In this genus there is characteristically a small white ear tuft. 
See Table 12 for an indication of size in this squirrel. 

Systematic history.— J. A. Allen (1906, p. 475) proposed the generic 
name of Tamiops, describing characters of the molariform teeth which 
differ from those of Sciurus (a genus which then included what are 
now the species of Callosciurus) . Pocock's (1923) evidence is rather 
equivocal on this matter of generic distinction of Tamiops, for al- 
though his example of the baculum in Tamiops differs sharply from 
those of some species of Callosciurus, it nevertheless closely resem- 
bles the example of lokroides. However, Robinson and Kloss (1918, 
p. 239) had already seized upon the generic separation of Tamiops 
and placed Tomeutes and six other genera between Callosciurus and 
Tamiops in their "A nominal list of the Sciuridae of the Oriental Re- 
gion ..." This placement, although unsupported by further evi- 
dence, has nevertheless been followed by Osgood (1932) and G. M. 
Allen (1940) in their extensive faunistic reports. It is especially 






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Fig. 17. Skull and left mandible of the striped tree squirrel of the Indochinese 
Subregion, Tamiops, AMNH No. 111396 of species smnhoei, Xl. 

noteworthy that although Chasen (1940) is too conservative to rec- 
ognize the genus Callosciurus (as distinct from Sciurus) in his "Hand- 
list of Malaysian mammals," he does recognize Tamiops as a full 
genus and places four other genera between it and "Sciurus." Zahn 
(1942, p. 76) separated Tamiops further still from Callosciurus when 
he sought to make Tamiops a subgenus of Funambulus. 

At the conservative extreme on the treatment of Tamiops Eller- 
man (1940, p. 347) accepted Tamiops as of subgeneric rank, and 
placed it in the genus Callosciurus. These treatments by Zahn, 
G. M. Allen, Ellerman, and Chasen are especially interesting since, 
being so contemporary, each presumably must have been offered 
without advantage of the others' thinking. Ellerman (1940, p. 350) 
points out that in describing Tamiops as a new genus J. A. Allen 
distinguished it only from species which are now in the restricted 
genus Sciurus and not from species of Callosciurus, and that diag- 
nostic characters attributed to Tamiops by Allen do not in fact dis- 
tinguish it from species of Callosciurus. In Simpson's (1945, p. 79) 
classification, then, Tamiops is included in Callosciurus. In a later 
classification of the Sciurinae (Moore, 1959, p. 198) the matter of 
subgeneric versus generic status for Tamiops was not reconsidered in 
any detail, and Tamiops was left as a subgenus of Callosciurus. Zahn 
(1942) in classifying Tamiops as a subgenus of Funambulus, ignored 
the evidence of their great difference shown by Pocock (1923) in the 
character of their respective bacula. On the basis of characters of 
the skull, also, Moore (1959, pp. 170, 173) has subsequently shown 
that Funambulus and Tamiops belong in separate tribes with the 
affinities Pocock had earlier attributed to them. Even in the pelage 
characteristics, which is Zahn's line of evidence for associating the 


two, the dorsal pelage color pattern of stripes differs in several funda- 
mental ways: (1) The midstripe and two additional pairs of stripes 
which are the same color as the midstripe, are the white ones in 
Funambulus, but the black ones in Tamiops. (2) The color of the 
pelage in between these stripes appears to be a rather uniformly 
colored area in Funambulus whereas in Tamiops it appears to be two 
pairs of stripes, the inner pair usually differing markedly in color 
from the outer pair. (3) The predominating stripes in Funambulus 
are the five including the middle one; in Tamiops the predominating 
stripes are the outer pair of the four interspaced between those five, 
and secondarily, the middle one of the five. 

Diagnosis. — The recent recognition of the distinctiveness of the 
genus Sundasciurus (Moore, 1958) constituted by several species 
which until then had been thought to belong in the genus Callo- 
sciurus eliminates some of the confusion and difficulty formerly found 
in attempting to distinguish the species of Tamiops from those of 
Callosciurus on characteristics of the skull. In addition to the light 
stripes of the dorsal pelage, which so strongly distinguish Tamiops 
from all other Callosciurini that earlier authors have been predis- 
posed to recognize Tamiops as a genus, there are now several char- 
acters of the skull which distinguish the species of Tamiops from the 
species remaining in the genus Callosciurus: (1) Small size associated 
with relatively fairly short orbit distinguishes Tamiops from Callo- 
sciurus; i.e., an orbit length of less than 13 mm. is Tamiops, more 
than 13 mm. is Callosciurus. (2) The midlateral processes of the 
basioccipital are obsolescent in Tamiops but generally well-developed 
in Callosciurus. (3) The upper profile of the rostrum is flat in Tami- 
ops, but generally somewhat arched in Callosciurus. These skull 
characters may here be considered distinctive at the level of genus 
for a tree squirrel (see discussion of conservatism in tree squirrels 
by Moore, 1959, p. 194) phylum which is polytypic. 

Intra-generic relationships. — Bonhote (1900) published the first 
revision of what is now Tamiops, recognizing nine forms, all as mem- 
bers of a single species mcclellandi. Robinson and Kloss (1918) ack- 
nowledged 12 forms of mcclellandi, and admitted swinhoei as a distinct 
species. Osgood (1932) considered the whole group again and listed 
13 named forms of swinhoei as "entitled to some sort of recognition" 
and nine forms of mcclellandi as "most likely to have permanent rec- 
ognition." He tentatively recognized four species: mcclellandi, swin- 
hoei, maritimus, and monticolus. Subsequently G. M. Allen (1940) 
reduced monticolus to synonymy and maritimus to subspecies status. 


Seasonal pelage changes in Tamiops, particularly in the nontrop- 
ical parts of its range, have been found to be pronounced. It is 
therefore important to consider whether the specimens one exam- 
ines are summer or winter skins, particularly in the forms from China. 
Actually the pelage appears to be virtually as long in summer skins 
as in winter skins. 

The skulls in Tamiops in general exhibit great uniformity. The 
smallest skulls appear to be in mcclellandi, barbei, and rodolphei. 
Skulls of these forms are nearly equal in both length and breadth. 
The skulls of the species swinhoei are typically considerably larger. 
See Table 12. The nasals and teeth of the species swinhoei are in 
general also larger than in the remainder. The subspecies barhei and 
rodolphei seem to have exceptionally small teeth, leaving mcclellandi 
in an intermediate position in this respect. 

The notably small size of all four Tamiops species (as shown in 
Table 12), the surprising narrowness of its tail for a tree squirrel, 
the flat dorsal profile of the rostrum, and the three pairs of functional 
mammae instead of two pairs — all suggest that the original Tamiops 
species could most reasonably have come from a stock in the Ma- 
laysian Subregion which also gave rise to the genus Sundasciurus, 
the smaller species of which may have outcompeted and replaced the 
Tamiops line there after the latter had successfully spread to the 

Tamiops mcclellandi (Horsfield) 

Definition. — The species Tamiops mcclellandi is constituted by 
subspecies mcclellandi, collinns, kongensis, barbei, inconstans, and leu- 
cotis, and see their synonyms. The species distribution as known 
from material examined, is shown in Figure 18. 

Diagnosis. — Species mcclellandi is generally smaller and more viv- 
idly striped than species swinhoei and has shorter, thinner pelage. 
The inner pair of light stripes is not as bright as the outer pair in 
mcclellandi, and the black middle stripe is not partially bisected 
longitudinally as is usual in rodolphei. 

Relationships to other species. — The species mcclellandi is princi- 
pally known from west of the Mekong River. Its relationships to 
the larger, more montane species swinhoei are described below in the 
account of the species swinhoei. The geographic relationship of spe- 
cies mcclellandi to species rodolphei is shown in Figure 18. These 
two species are ecological equivalents as far as is known. The species 
mcclellandi appears to reach across the Salween and Mekong rivers 




A^' ^ 



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30°U + 


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Fig. 18. Ranges of the Burmese striped tree squirrel, Tamiops mcclellandi, 
A to F; and the Cambodian striped tree squirrel, Tamiops rodolphei, G and H, 
plotted from collecting localities of material examined. Subspecies of mcclellandi: 
A, mcclellandi; B, collinus; C, kongensis; D, barbei; E, leucotis; and F, inconstans. 
Subspecies of rodolphei: G, rodolphei; and H, elbeli. 



and through Yunnan to northern Vietnam (where it is represented 
by the subspecies inconstans) . There it seems to be sympatric with, 
or interdigitated with, species maritimus. 

Intraspecific variation. — The species mcclellandi occurs in forests 
from Sikkim, Bhutan, Assam, and northernmost Burma southward 
in forested hill country probably throughout Burma, Thailand ex- 
cepting the Mekong plain, and the Malay States. Its outer pair of 
light stripes in the dorsal pelage is strikingly light and prominent in 
comparison to the subdued inner pair, and it is characteristic that 
the outer light stripes continue strongly marked across the shoulder 
area and connect with the light facial stripe. 

While five subspecies of mcclellandi receive recognition here, the 
authors wish to point out that there is a greater distinction between 
the form mcclellandi and the other four than there is between any 
two of the other four. Because of this, one of the present authors 
(Tate) elected to recognize mcclellandi as a monotypic species and 
the other four forms as constituting the species barbei. This may 
prove to be correct if study of further material discloses no intergra- 
dation between mcclellandi and collinus. One might suppose it to 
be equally logical in our present state of knowledge, however, to con- 
sider the distinction between mcclellandi and the other four forms as 
of subspecific value and that the "other four forms" constitute a vari- 
able but single subspecies. From the material that we have studied, 
it seems to us that leucotis, barbei, and kongensis are distinct geo- 
graphic subspecies occupying equally distinct physiographic areas. 
The other form, collinus, is somewhat less distinct, and also the 
considerable geographic area it occupies does not appear to be physio- 
graphically distinct. However, this form is more than a broad zone 
of intergradation between mcclellandi and kongensis, for it possesses 
a color character shared by neither, the blackness of the inner paired 
dark lines. It is here considered, therefore, that the distinction be- 
tween subspecies mcclellandi and the other four is at an advanced 
subspecific (and perhaps incipient species) state of differentiation; 
whereas the distinctions between any adjacent two of the subspecies 
collinus, kongensis, barbei, and leu/^otis are at a subspecies, but less 
advanced, state of differentiation. 

Tamiops mcclellandi mcclellandi (Horsfield) 

Sciurus mcclellandi Horsfield, 1839, Proc. Zool. Soc. London, 1839, p. 152. 
Sciurus pembertoni Blyth, 1842, J. Asiatic Soc. Bengal, 11, p. 887. 
Sciurus macclellandi manipurensis Bonhote, 1900, Ann. Mag. Nat. Hist. (ser. 7), 
5, p. 51. 


Types. — Sciurus mcclellandi, two cotypes, BM Nos. 
373, adult male and a young adult, the former marked "lectopara- 
type," both from "Assam"; pembertoni, BM No., adult 
female, from Bhotan; manipurensis, BM No., old male 
from Aimole, Manipur. 

Material examined, from Assam, India. — Tura, Garo Hills 
(AMNH), one; Tura Mt., Garo Hills (CNHM), three; Pynursla, 
Khasi Hills (CNHM), two; Mawphlang, Khasi Hills (CNHM), 
three; Cherrapunji (CNHM), one; Sangau, Lushai Hills (CNHM), 
three; Karong, Manipur (CNHM), six; Takubama, Naga Hills 
(CNHM), four; Kohima, Naga Hills (CNHM), two; Sadiya, Mish- 
mi Hills (AMNH), three, (CNHM), one; Dreyi, Mishmi Hills 
(AMNH), one; Tezu, Mishmi Hills (AMNH), five. 

Material examined, from Sikkim, India. — Darjeeling (CNHM), 
one; Rungbong Valley, 4000 feet (MCZ), one; Chungtang (CNHM), 
three, (USNM), one; Lingtam (USNM), three, (CNHM), 28; "Sik- 
kim" (MNHN), one. 

Material examined, from Burma. — 6 to 25 miles N. of Myitkyina 
(USNM), five; 170 miles N. of Myitkyina (CNHM), one; Seniku- 
Shingaw Road (AMNH), one; Laukhaung-Pyepat Road (AMNH), 
one; Nanyaseik (AMNH), three; Lonkin (AMNH), one; Mansum 
(AMNH), one; Dalu (AMNH), four; Tagahka (AMNH), five; Luck- 
changa (AMNH), one; Dagung Hka (AMNH), one; Haibum 
(AMNH), three; Singkaling Hkamti (AMNH), one; Kawai, W. Bank 
(AMNH), one; Kaunghein (AMNH), four; Nonswa (AMNH), one; 
Mawlaik (AMNH), one; Mt. Victoria, Pakokku Chin Hills (AMNH), 

Type description. — Typically mcclellandi, studied in 1951, has the 
median black line about 4 mm. wide and extending from nape to 
rump. The lateral pair of dark lines extends from the shoulders to 
the rump but are a little wider, 5 mm. They are partly obscured at 
their inner edges by the overlying yellowish buffy hair tips of the 
intervening area. Laterally they are edged by the pair of strongly 
colored yellow-buff lines which are the primary elements in the dor- 
sal pattern. Then two yellow lines reach from the rostrum over the 
whiskerpatch, beneath the eyes and ears, become broadened at the 
sides of the neck and connect with the outer pair of lines on the back, 
which finally fade out at the rump. The maximum width of this line 
is at the side of the neck. External to this pair of yellow lines is the 
merest suggestion of the outermost pair of dark lines about 1 inch in 


length and less than 2 mm. in width. Underparts buffy-tipped (with- 
out any trace of red) with the bases of the hairs gray. Hands and feet 
grayish buff. Tail with bases of the hairs brownish buff, subterminal 
bands black, tips buffy white. Extreme tip of tail blackish. This is 
the basic form against which all others must be compared. The 
length of the head and body is ca. 110 mm., of tail ca. 100 mm., hind 
foot ca. 28 mm. It is apparently a much smaller Tamiops than any 
member of the species swinhoei. 

Discussion. — While there are slight differences in pelage color 
which may be locally geographic in expression within this typical 
subspecies, these are negligible by comparison with the differences 
between T. m. mcclellandi and its nearest conspecific neighbor T. m. 
collinus. A series collected high on Mt. Victoria in Burma is not 
only lighter in ventral pelage color than typical mcclellandi but some- 
what larger in size. Nevertheless, these differences would make of it 
only some sort of a micro-race by comparison with those differen- 
tiating collinus and mcclellandi. 

Habits. — In a letter of June 10, 1961, Lord Cranbrook contributes 
the following field notes on this species from his collections on the 
Adung Valley expedition: "No. 60. The commonest squirrel in the 
district (Myitkyina — Sumpra Bum). Usually seen high up in tall 
trees, moving in short rushes and then staying motionless, sometimes 
head downward, often some minutes at a time. I have never seen 
one in low bushes, always high up. The tail is usually held straight 
out [behind] and very seldom 'flicked' or held over the back. Fright- 
ened, they freeze flattened out with tail extended." These notes were 
not included in Cranbrook's account in Kinnear (1934). 

Another collector wrote: "This squirrel is found in most of the 
forests above 5,000 feet. It is common but owing to its power of 
concealment is generally not very easy to find. It is very nimble and 
appears to glide rather than walk along the boughs and slender twigs. 
I observed it in pairs and sometimes in small parties often sharing 
the tree with D. lokriah. It seldom comes to the ground. The call 
is a quickly repeated 'chick' much harsher in tone than that of 
F. pennanti." (C. A. Crump in Wroughton, 1916a, p. 488.) 

Tamiops mcclellandi collinus Moore 

Tamiops mcclellandi collinus Moore, 1958, American Mus. Novitates, no. 1879, 
p. 1. 

Type. — AMNH no. 163528, an old male from 800 meters elevation, 
Maymyo, Burma, collected November 28, 1937, by Gerd Heinrich. 


Material examined. — Maymyo, 800 meters, Burma (AMNH), 14; 
Mengting, 1700 feet, Yunnan (AMNH), four; Mu-cheng, 5000 feet, 
Yunnan (AMNH), two; Nam Ting, Yunnan (MCZ), one; Chiang 
Saen Kao, Chiang Rai, Thailand (USNM), one; Sawan Mt., Ban 
Seid, Loey, Thailand (USNM), one; Lomloe Mt., Maeo, Goksatawn, 
Loey, Thailand (USNM), eight; Vientiane, Laos (USNM), five; 
Chieng Dao, Thailand (CNHM), one; Doi Chieng Dao, 4000 feet, 
N. Siam (ANSP), three; Chieng Sen [Chiang Saen], N. Siam (ANSP), 
one; Don Qua, Laos (ANSP), one. 

This subspecies is more strongly striped than mcclellandi, less red 
than barhei, darker dorsally than kongensis, and much more strongly 
striped than inconstans. It is not in geographic contact with leu- 
cotis, but is ventrally more orange and less yellow than leucotis. 

Tamiops mcclellandi inconstans (Thomas) 

Tamiops inconstans Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 306. 

Type. — BM No., an old male from southern Yunnan 
(possibly near Mengtsze), collected January 31, 1910, by H. Orii. 

Material examined.— "Yunnsin" (BM), one; "Tonkin" (MCZ), 
three, (CNHM), one; Bao Ha, Tonkin (CNHM), two; Pakha, An- 
nam [Tonkin] (CNHM), one. 

Original description. — "This very distinct little squirrel is char- 
acterized by its unusually inconspicuous striping above and by the 
strong yellowish buff y of its lower surface — in fact, it is above one of 
the dullest and below one of the brightest of the genus." We find 
the ventral pelage to be Light Salmon Orange where the colored hair 
tips entirely cover the gray hair bases. 

Tamiops mcclellandi kongensis (Bonhote) 

Sciurus mcclellandi kongensis Bonhote, 1901, Proc. Zool. Soc. London, 1901 
p. 55. 

Type. — BM No., adult female from Raheng, Siam, col- 
lected February 7, 1900, by T. H. Lyle. 

Material examined, all from Thailand.- — Chieng Dao [Ban Chiang 
Dao] (USNM), one, (CNHM), one; Mae Hong Sorn [Ban Muai To] 
(USNM), two; Doi Hua Mot (USNM), one; Doi Phra Chao [Khao 
Pha Cho] (USNM), one; Mae Wan River near Doi Saket (MCZ), 
one; Nan [Muang Nan] (USNM), four; Doi Sutep [Doi Suthep] 
(AMNH), one; Doi Angka [Doi Ang Ka=Doi Inthanon] (MCZ), 
six, (USNM), two; Koon Tan [Doi Khun Tan or Sathani Khun Tan] 


(USNM), four, (ANSP), two, (NR), three, (CNHM), one; Chum 
Poo [Sathani Tha Chomphu] (NR), two; Mesarieng [Ban Mae Sari- 
ang] (USNM), one; Ta Chang Tai [Tha Chang Tai] (USNM), two; 
Raheng [Ban Rahaeng] (USNM), five; Chiengmai [Muang Chiang 
Mai] (ANSP), two, (MCZ), three; "Wung Pratart Farm" Kam Peng 
Pet Prov. [Kampheng Phet] (CNHM), 14; 40 miles E. of Um Pang 
[Ban Um Phang] (AMNH), one; Pitsanulok (AMNH), one; Pak Tha 
(NR), one; Pak Koh (NR), one; "Den Chai" (NR), one; Pran [Pran 
Buri] (USNM), two; Huey Yang, Sriracha [Sathani Huai Yang] 
(USNM), one; "Muang Qua Yie," Nakorn Sawan (USNM), one; 
"Wat Sytie," Paknampho, Nakorn Sawan (USNM), one; "Ban 
End," Chiengmai (USNM), one; Pujeg, Pak Tho, Rajburi (USNM), 
two; Kowjeen, Pak Tho, Rajburi (USNM), one; Tungnockarien, 
Jombing, Rajburi (USNM), three; Borploy, Kanjanaburi (USNM), 
two; "Tarn bol, Tak tog, Bantak," Tak [Ban Rahaeng] (USNM), 
two; Tra Khanun, Hin Laem, Kanachanaburi (USNM), nine; Ban 
Klua Klan [Ban Khlua Klang], Prachuap Kiri Khan (USNM), one; 
Klong Klung [Ban Khlong Khlung], Kampheng Phet (USNM), one; 
"Ban Thoong Cheark, Slog Bartara, Kha Nu," Kamphengphet 
(USNM), two; "Srisawat" (ANSP), two. 

In this race the dimensions of the various dorsal lines remain al- 
most exactly as in harhei. The chief difference occurs in the overall 
pallid coloration. There is a general infusion of light gray, instead 
of the reddish orange wash of barbei and the yellow wash in leucotis. 
The pale hues of the back in the present race are very pale buffy 
yellow, and the underparts are about Ochraceous Buff. This sub- 
species seems to have differentiated in the rain shadow zone of Thai- 
land, and has a range apparently identical with that of the pale race 
of Menetes herdmorei consular is. 

Tamiops mcclellandi barbei (Blyth) 

Sciurus barbei Blyth, 1847, J. Asiatic Soc. Bengal, 16, pt. 2, p. 875. 

Type. — Not seen. From Zami River, Ye Province, 100 miles 
south of Moulmein, Tenasserim, Burma. 

Material examined, all from Burma. — Kawkereik, Tenasserim 
(AMNH), two; Lampha, Tenasserim (AMNH), one; Tavoy 
(CNHM), one; Tenasserim Town (CNHM), one; Taok Village, 
Tenasserim (AMNH), two; Zami River, Ye Province, 100 miles 
south of Moulmein (BM), one. 


Discussion. — While material from Kawkereik at 175 feet eleva- 
tion has ventral pelage about Ochraceous-Salmon at the throat and 
Cinnamon-Rufous or Ferruginous on the remainder of the venter, 
material from nearby Taok at 1100 feet elevation is much paler ven- 
trally and slightly grayer dorsally so as to be really intermediate 
between barhei and kongensis. Like the Taok material, also, is a 
specimen from neighboring Lampha. Ventral pelage of these is near- 
est to Capucine Orange. The Kawkereik and Lampha material was 
collected December 12 and 28 respectively, the Taok January 16. 
The entire dorsal pelage of the Kawkereik material is infused with 
reddish which shows as Light Salmon Orange in the outer light dor- 
sal stripes. 

Habits. — Field observations are offered from southernmost Burma: 
"Completely arboreal. I have never observed this species actually 
on the ground. Common around villages on the Tenasserim River 
but not observed further south. More plentiful in native fruit gar- 
dens than in jungle. Weight, 13^-2 ozs." (G. C. Shortridge in 
Wroughton, 1915b, p. 713.) 

Tamiops mcclellandi leucotis (Temminck) [Extraterritorial] 

Tamias [sic] leucotis Temminck, 1853, Esquisses Zool. Cote Guine, p. 252. 
Sciurus novemlineatus Miller, 1903, Proc. Biol. Soc. Washington, 16, p. 147. 

Types. — Tamias leucotis "Mallaca peninsula" and not seen by 
the present authors; novemlineatus, USNM No. 84403, adult male 
from Trong, 1500 feet, collected February 19, 1897, by W. L. Abbott. 

Material examined, from Malay States. — "Selangor" (USNM), 
two; Ginking B., Selangor (USNM), one; Brinchong Hill, 5500 feet, 
Cameron Highlands, Pahang (USNM), one; Gunong Mengkuang, 
4800 feet, Lebah, Selangor (USNM), one; Telom, 3800 feet, Perak 
near the Pahang border (USNM), one. 

Material examined, from peninsular Thailand. — Trong [Trang], 
500 feet (USNM), one; Khaw Nom Phu, 2000 feet, Trong [Trang] 
(USNM), one; Ban Kiriwong (USNM), five; Ban Chit [Ban Nong 
Chik] (USNM), one. 

Material examined, from Tenasserim, Burma. — Telok Besar 
(USNM), three. 

Discussion. — This subspecies lacks the general reddish infusion in 
the dorsal pelage displayed by T. b. barbei, and the ventral pelage 
of leucotis is much the yellower, being Ochraceous Buff in the yellow- 


est instances and a little more orange than that in several others, 
matching no Ridgway color precisely. Surprisingly the series of 
three from Trong show strong evidence of intergradation with barbei 
both dorsally and ventrally, the ventral pelage being Apricot Buff 
on the throat and Orange Cinnamon on the venter. Yet from north 
of this, 200 miles closer to barbei range, at Telok Besar in the south- 
ern tip of Burma, the specimens show no dorsal infusion of red, and 
the ventral pelage is also correct for leiLCotis. Presumably less than 
100 miles north of this leucotis intergrades with both barbei and 

Habits. —Concerning the distribution and habits of this subspecies 
Robinson {in Bonhote, 1903, p. 21) wrote, "This beautiful little spe- 
cies was not met with on the Eastern side of the Peninsula. In Perak 
and Selangor it is certainly a mountain form, and I do not think it 
occurs much below 3000'. It was very common at Telom, and was 
also very abundant on the mountains round the Semangko Pass. 
It is very largely an insectivorous species, and seems to keep chiefly 
to the trunk and main branches of the trees, running along them 
with its tail pressed close against the bark." Chasen (1940, p. x) in 
commenting on the distinctive mountain fauna encountered above 
3000 feet elevation in the Malaysian Subregion, notes that, "A 
Himalayan element (e.g., Rattus bowersi, Tamiops, Dremomys rufi- 
genis, Sciurus erythraeus) is the dominant feature of the high-levels 
in Perak, Pahang and Selangor, but it fades out in the south, and is 
absent in Negri Sembilan and on the Johore hills; it is also weak on 
the isolated summit of Kedah Peak in the north." 

Tamiops rodolphei (Milne-Edwards) 

Definition. — The species rodolphei is constituted by subspecies 
rodolphei and elbeli and the named forms here included in them. 
Its distribution, shown in Figure 18, is in Cambodia, southern Laos, 
southern Vietnam, and eastern Thailand. 

Diagnosis. — Species rodolphei is distinguished from the other spe- 
cies of this genus by: (1) The outer and inner pairs of light lines are 
of equal width. (2) The inner pair of light lines approximates the 
brightness of the outer pair. (3) The blackish middorsal line is 
almost invariably divided for part of its length by a very thin, pale 
brown line. 

Relationships to other species. — Tamiops rodolphei appears to re- 
place Tamiops mcclellandi with no known overlap or intergradation 
in eastern Thailand. In southern Vietnam it overlaps or interdigi- 


tates with the known southern extremity of the range of the species 
maritimus. Compare Figures 18 and 20. 

Tamiops rodolphei rodolphei (Milne-Edwards) 

Sciurus (Tamias) rodolphei Milne-Edwards, 1867, Revue Mag. Zool., 19, 
p. 227. 

Tamiops macclellandi liantis KIoss, 1919, J. Nat. Hist. Soc. Siam, 3, p. 370. 

Tamiops lylei Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 307 (hom- 

Tamiops macclellandi dolphoides Kloss, 1922, J. Nat. Hist. Soc. Siam, 4, p. 101. 

Callosciurus holti Ellerman, 1940, Families and Genera of Living Rodents, 1, 
p. 355 (new name for lylei). 

Types.— Sciurus (Tamias) rodolphei MNHN No. 1864-683 (318), 
adult from Saigon, Indo-China; dolphoides, apparently lost, Kloss 
field no. 2723/CBK, adult female from Kompong Som Bon, near Sr4 
Umbel, southwest Cambodia; liantis, USNM No. 221542, young 
adult female from Sata Hip, near Cape Liant, southeastern coast of 
Siam; lylei= holti, EM No., young adult male from sea coast 
fifty miles south of Bangkok, Siam. 

Material examined, from Thailand. — Sriracha [Ban Si Racha] 
(USNM), two, (CNHM), one; Chantabun [Chanthaburi] (ANSP), 
two, (USNM), one; Kao Sabab [Khao Sa Bap] (USNM), two; Lem 
Ngop [Ban Laem Ngop] (USNM), one; Kao Seming, Krat [Khao 
Saming, Trat] (ANSP), two, (USNM), one; Nongkhor [Ban Nong 
Kho] Chon Buri Prov. (USNM), two. 

Material examined, from Indochina. — Ban Me Thuot [Buon Ma 
Thuot], Annam (CNHM), four; Bien Hoa, Cochin China (USNM), 
two; Gougah [Lien Gongah] Annam (CNHM), 11, (MCZ), one; 
Thateng (USNM), one, (CNHM), 11; Pak-S6, Laos (CNHM), two; 
Paksong, Laos (CNHM), one; "Phu Kongutoul" (CNHM), one; 
Quangtri Phuoc Mon, Annam (CNHM), three; Ninh Hoa, Annam 
(CNHM), one; "Cochin China" (CNHM), one; Plateau Bolovens, 
2500-3000 feet, Laos (ANSP), three, (AMNH), three; Tayninh, 100 
feet, Cochin-China (MNHN), one; Angkor, 200 feet, Cambodia 
(MNHN), two; N. V. Kampot, 100 feet, Cambodia (MNHN), one. 

The head and nape are almost uniformly dull brown in this sub- 
species. The tail is exceptionally slender due to shortness of its 
hairs. The underparts tend toward yellow or orange, somewhat as 
in mcclellandi kongensis or m. collinus, never buffy or whitish as in 
m. mcclellandi and in subspecies of maritimus. 


Tamiops rodolphei elbeli Moore 

Tamiops rodolphei elbeli Moore, 1958, Amer. Mus. Novitates, no. 1879, p. 4. 

Type.—VSNM No. 294876, adult female, skin and skull, collected 
by Robert E. Elbel on January 18, 1952, at village of Ban Lad, 
Pookeio District, Chaiyaphum Province, Thailand (ca. latitude 
16° 40' N., longitude 101° 45' E.). 

Material examined. — Ban Lad, Pookeio, Chaiyaphum, Thailand 
(USNM), two; Ban Non Toulek, Pookeio, Chaiyaphum, Thai- 
land (USNM), three; Khon Kaen, Thailand (USNM), seven; 
"Watpa" (ANSP), one. 

Diagnosis. — Tamiops rodolphei elbeli is distinguished from all sub- 
species of Tamiops mcclellandi, swinhoei, or maritimus by the mid- 
dorsal black stripe, for this stripe is divided at least part of its length 
by a fine line of yellowish brown down its middle. It differs from 
Tamiops rodolphei rodolphei, which occurs to the east and south of it, 
in the brilliance of its Isabella Color crown and nape compared with 
the Buffy Brown (XL) to Snuff Brown (XXIX) crown and nape of 
T. r. rodolphei (including three virtual topo types from Bien Hoa, 
Cochin China). 

Dimensions. — The skull measurements (in millimeters) of the 
type are: occipitonasal length, 32.6; condylobasal length, 30.0; right 
maxillary toothrow, 5.4; palatal length, 15.8; mastoid breadth, 14.9; 
greatest breadth, 20.1; nasal length, 10.2; and orbitonasal length, 

Field measurements in millimeters of the total length, tail length, 
and hind foot of the type of T. r. elbeli are, respectively, 250, 130, 
and 30. The same characters recorded for a male and two females 
of T. r. elbeli from Ban Non Toulek are, respectively: 220, 250, 220; 
110, 129, 102; and 30, 32, 30. 

Variation. — Of 17 skulls of Tamiops m. collinus and kongensis 
examined in the United States National Museum from localities geo- 
graphically near to elbeli, five have the posterior surface of the ex- 
tremities of the upper incisors beveled so that it shows just a slight 
concavity in the side view. Each of the other 12 is cut so that it 
shows a notch. Of 13 skulls of T. r. elbeli examined, four have just 
a slight concavity, and nine have a straight bevel. The lateral mar- 
gins of the basioccipital are curled ventrad and rise slightly to a low 
point or process in T. r. elbeli, but in T. m. collinus and kongensis 
and also T. r. rodolphei these edges remain low ridges and do not 
form a point or process. 


Discussion. — This subspecies comes from the eastern part of Thai- 
land, the Mekong plateau 2000 feet in elevation, which is subtended 
on the north and east by the Mekong River and south by the Phanom 
Dang Raek, a range of mountains. Excepting for Gyldenstolpe (1914) 
who collected mammals on its southwestern edge about Korat, evi- 
dently no mammalogist had collected on this plateau before R. E. 
Elbel got into the northwestern quarter of it. The material repre- 
senting Tamiops r. elbeli all comes from Chaiyaphum Province and 
Khonkaen Province. 

Tamiops swinhoei (Milne-Edwards) 

Definition. — This species includes subspecies swinhoei and the 
several named forms here included in it, and subspecies vestitus. 
Figure 19 shows the distribution of species swinhoei as known to us 
from specimens. 

Diagnosis. — This is apparently an extremely variable species geo- 
graphically, and although swinhoei seems to be generally larger than 
the other species of Tamiops and to have longer, denser fur, diag- 
nostic distinction by these characters does not seem possible in the 
material we have examined. These two tendencies may be related 
to the fact that species swinhoei lives farther north or at higher ele- 
vations than the other species of Tamiops. The outer pair of light 
lines is less brilliant than those of mcclellandi and often broader, and 
they more generally stop at the shoulder instead of being connected 
with the cheek stripe. 

Relationships to other species. — The range of this species appar- 
ently does not overlap that of mcclellandi to any great extent. The 
only localities from which both species have been taken are where 
the edges of their ranges meet. At these places there has been a 
notable difference in the elevations at which the two species were 
taken. In northeastern Burma near Imaw Bum Anthony (1941, 
p. 93) took the species mcclellandi between 1000 and 2000 feet ele- 
vation in definitely tropical habitat and at 4500 feet, but he collected 
the species swinhoei at 9000 and 10,000 feet. Northward about 130 
miles from Imaw Bum, Cranbrook collected mcclellandi at 1500 feet 
elevation but at nearby Adung Valley took swinhoei at 8000 and 9000 
feet. At Mucheng, Yunnan, China, about 175 miles south of Imaw 
Bum, Edmund Heller collected the species mcclellandi at 5000 feet 
elevation and swinhoei at 7000 feet. These three localities are the 
westernmost known for the species swinhoei and the northeastern- 
most known for mcclellandi. See Figure 19. 






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Fig. 19. Distribution of the Chinese striped tree squirrel, Tamiops sivinhoei 
(entire range) and the maritime striped tree squirrel, Tamiops maritimus (north- 
eastern range) as obtained from plotting collecting localities of materials examined. 
Subspecies of swinhoei: A, swinhoei; and B, vestitus. Subspecies of maritimus: 
C, maritimus; and D, mx)nticolus. 

On the apparent southern edge of the range of T. swinhoei Dela- 
cour and Lowe evidently obtained a series of 17 from 8000 to 10,000 
feet elevation on Mt. Fan Si Pan near Chapa in northern Tonkin, 
but at lower elevations in the same vicinity took T. maritimus haina- 
nus. In reporting this, Osgood (1932, p. 292) recognized these ex- 
amples of swinhoei as a race of monticolus. Eastward in the range of 
swinhoei there is an extraordinary gap in examined material of this 
species, as shown on the map. The eastern distribution (T. s. ves- 
titus) lies entirely in Wang's (1956) oak-dominated deciduous broad- 
leaved forest type. The area of this forest type extends westwardly 
about to the western distribution (T. s. swinhoei), but it appears that 
not a single one of the twenty-odd localities that we have for T. swin- 
hoei here is in that general forest type area. Possibly this constitutes 


something of a barrier, and the absence of records of Tamiops from 
middle China really indicates absence of the genus, for Sciurotamias 
has been collected there. Compare Figures 19 and 33. Of course, 
Sciurotamias is very much a ground squirrel and would possibly have 
increased in the same places where Tamiops may have been elimi- 
nated by deforestation. Fu (1936) found "Tamiops macclellandi 
vestitus" in the Sung-shan, which he describes as a famous mountain 
in western Honan with luxuriant forests, and he says that this squir- 
rel lives in holes in trees. Other than this we find no literature 
references to Tamiops swinhoei in very central China. 

Tamiops swinhoei swinhoei (Milne-Edwards) 

Sciurus maccleUandi var. swinhoei Milne-Edwards, 1874, Recherches . . . des 

mammiferes, 1, p. 308. 
Tamiops clarkei Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 304. 
Tamiops maritimus forresti Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, 

p. 305. 
Tamiops spencei Thomas, 1921, J. Bombay Nat. Hist. Soc, 27, p. 503. 
Tamiops macclellandi russeolns Jacobi, 1923, Abh. Berich. Mus. Dresden, 16, 

no. 1, p. 11. 
Tamiops monticolus olivacetLS Osgood, 1932, Field Mus. Nat. Hist. Publ., Zool. 

Ser., 18, p. 292. 

Types. — Sciurus maccleUandi var. swinhoei, type not found in the 
MNHN in Paris, but there are four possible cotypes taken at Moupin 
by Pere David in 1870, MNHN 1870-608 (309); 1870-39 (311); 
1870-38; and 1870^0 (308); russeolus, three cotypes (skins without 
skulls) from Tsalila, a pass near Atuntse, between the upper Yangtse 
and upper Mekong Rivers, SMTB 5887-9 collected September, 1916- 
1917 [1914] by Hugo Weigold; clarkei, BM No., adult male 
from forests in Yangtse Valley, latitude 27° 20' north, 8000 feet col- 
lected September, 1918, by George Forrest; forresti, BM No. 20.1.- 
16.4, adult male from Lichiang Range, Yunnan, 27° 20' latitude 
north, 11,000 feet collected July, 1918 by George Forrest; spencei, 
BM No., female from 28° 22' north latitude, 97° 40' east 
longitude, at 10,000 feet; olivaceous, BM No., adult male 
from Lo Gui Ho, Tonkin, 5000 feet, collected December 30, 1916 by 
P. M. Leonard. 

Material examined, from Tibet. — Tsarong, 7000 feet, Londjre 
Valley (USNM), one. 

Material examined, from Sikang, China. — Mupin [Muping] 
(MNHN), one, (USNM), two; Luting Shan [Lutingkiao] (CNHM), 


one; Wuchi [Wushi] (CNHM), one; Kulu (CNHM), one; "Ta 
Chiao," 12,000 feet near Tatsienlu (MCZ), one; 35 miles east of 
Hokow (ANSP), one; Madischung Valley, 10 miles from Hokow 
(ANSP), one. 

Material examined, from Kansu, China. — Chulungapu, 8000 feet, 
Upper Tebbuland (CNHM), one, (MCZ), six. 

Material examined, from Szechwan, China. — "Lian-feng-Kiang," 
Omei Shan (CNHM), one; 25 miles west of Wench wan, 5-9000 feet 
(AMNH), nine, (USNM), three; Wa Shan (USNM), one; Weichow 
[Weikiu] (USNM), one; "Tsungshi (Ngoloshi)" (ANSP), one, 
(MCZ), one. 

Material examined, from Burma. — Adung Valley, 8-9000 feet, 
Kachin Prov. (CNHM), three; Imaw Bum (AMNH), two; Road to 
Chimeli Pass, 10,000 feet (AMNH), two. 

Material examined, from Yunnan, China. — "Mt. Djinaloko," 
Yangtze Valley slopes, 10,000 feet (USNM), one; "Hofuping Mts.," 
Mekong Valley (USNM), eight; "Sila Mts.," Salween-Mekong Di- 
vide, 8000 feet (USNM), one; N. slopes of Likiang Snow Range 
(USNM), one; Likiang, 10,000 feet (AMNH), three; 40 miles N. of 
Likiang (MCZ), one; "Tao Mung Chung" southwest of Lu-tien 
(MCZ), one; Mu Cheng, 7000 feet (AMNH), three, (MCZ), one; 
Ngu-luko (CNHM), one; "Yunnan" (CNHM), two. 

Material examined, from Indochina. — Lo-Gui-Ho, Tonkin 
(CNHM), three; Chapa, Tonkin (CNHM), five. 

Pelage color. — Compared with mcclellandi, T. s. swinhoei is often 
distinguished by its much larger size and long, soft pelage. The 
general dorsal color is yellowish brown. The black middorsal line 
is much wider than in mcclellandi, 9 to 10 mm. The width of the 
pair of black lateral lines is ca. 7 mm., and their length ca. 75 mm. 
The outer light line is in swinhoei dull yellow brown (scarcely brighter 
than the yellow brown lines on either side of the median line) . This 
line is interrupted at the shoulder, not connecting with the cheek 
line which extends from beneath the ear forward to the whisker 
patch. The outermost dorsal line is black. Its dimensions are about 
28 by 5 mm. The underparts are buffy white with the bases of the 
hairs slate gray. 

Discussion. — A circle only 100 miles in radius will include the 
type localities of russeolus, clarkei, forresti, and spencei, and one of 
150 miles radius will include the type localities of these four and that 
of swinhoei as well. These localities are on high elevations in the 


mountains segregated by the deep, north-south river gorges which 
dissect the general area. Two of the type locaHties, furthermore, 
those of russeolus and forresti, are not even so separated from each 
other. We cannot concur in G. M. Allen's (1940, p. 669) acceptance 
of clarkei as a good subspecies. His distribution map does not in- 
clude in the range of T. s. swinhoei the Yunnan locality of Mucheng, 
nor in the range of clarkei the Yunnan locality of Taomungchung 
southwest of Lutien, although he identified material from these 
places respectively as T. s. swinhoei and T. s. clarkei. The inclu- 
sion of Mucheng brings the range of swinhoei 300 miles farther south 
than he had it shown, and the inclusion of Taomungchung southwest 
of Lutien would extend his shown range of clarkei nearly 200 miles 
east. The specimen from Taomungchung seems to us as different 
from Likiang material as either is from a series of swinhoei from near 
Wenchwan, and that if it were represented by a series, it would have 
as good a claim on subspecific difference from typical swinhoei as 
does the material from Likiang. Another subspecific name for one 
specimen from one locality could be justifiably added to the list of 
synonyms by the next author, however, unless some further collect- 
ing is done in the range of this species to demonstrate the geographic 
relationships of this and some of the other considerable variation 
which is already represented by names in the presently available 

Habits. — In a letter of June 10, 1961 Lord Cranbrook contributed 
the following observations on this species from his field notes of 30 
years earlier in the Adung Valley of Burma: "No. 206. Shot in a 
tree amongst scrub jungle just below snow line (March). When 
first seen quite low down in bushes and low tree rhododendrons, very 
unlike the habits of the striped squirrels [T. mcclellandi] seen further 
south, which keep entirely to high trees. Tail held straight out be- 
hind, not flicked. No. 229. Shot running along a fence around a 
field." For other details see Cranbrook's account of the expedition 
in Kinnear (1934). 

Tamiops swinhoei vestitus (Miller) 

Tamiops vestitus Miller, 1915, Proc. Biol. Soc. Washington, 28, p. 115. 

Type.—USNM No. 199561, adult male from Hsin-lung-shan, 65 
miles northeast of Peking, collected February 15, 1915, by Arthur de 
Carle Sowerby. 

Material examined, all from Chihli [Anhwei] Province, China. — 
Tungling (AMNH), six, (CNHM), six, (MCZ), one; Eastern Tombs 


(AMNH), six, (MCZ), two; Hsing-lung-shan, 65 miles N.E. of Pe- 
king (USNM), 12; 80 miles E. of Peking (USNM), two. 

Pelage color. — This race seems to be a pallid offshoot of the moun- 
tain dwelling subspecies swinhoei. As is also found in Tamiops mari- 
timus of southern China, there is a striking difference between 
summer and winter pelage. This is exhibited, for example, by seven 
specimens taken in April while still in the winter pelage in which 
only the midstripe is black, and five specimens taken in late July 
and early August in the summer pelage which differs by the mid- 
stripe and inner dark pair of stripes all being black. The outer and 
inner pairs of light lines are whitish, the outer pair whiter than the 
inner. The underparts are buffy white, their hair bases gray. 

Discussion. — Jentink (1883) records two specimens of this sub- 
species in the Leiden Museum from Tingchow, which is apparently 
120 miles southwest of Peiping. It is also this form which Fu (1936) 
reported in the Sung Shan of western Honan. 

Tamiops maritimus (Bonhote) 

Definition. — This evident species includes subspecies maritimus, 
monticolus, hainanus, and moi, and see the synonyms of these. The 
known distributions of maritimus and monticolus are shown in Fig- 
ure 19, those of hainanus and moi in Figure 20. 

Diagnosis.— The pelage is comparatively short and thin, the gen- 
eral coloration tends to be more olive, and the inner pair of light 
stripes tends to be nearer the color of the nape, than in the other 
species of Tamiops. 

Relationships to other species. — This is a relatively low altitude 
species occupying the southeastern coastal region of China and all 
of Vietnam and Laos, and according to G. M. Allen (1940) appar- 
ently penetrating to middle China. At the southern end of its range 
it ascends the Langbian Peaks to considerable elevation, but it is 
evidently also found at the foot. At Pak Hin Bun, Laos, Robinson 
found it along the Mekong River and on islands in the River. Al- 
though we have examined specimens of it from two other Laos local- 
ities along the Mekong, Vientiane and the westernmost locality, Lo 
Tiao, it is not known from west of that river. Its geographic rela- 
tionships to Tamiops swinhoei east of the Mekong in China remain 
too much a mystery. It is at least of interest that Thomas (1920, 
p. 305) once thought that maritimus penetrated Yunnan to the Liki- 
ang Range. When it is discovered what the distributions of these 
striped squirrel species are in central and southern Yunnan, it should 



become possible to interpret with greater confidence the information 
already available from northern Yunnan and nearby Burma, Sikang, 
and Szechwan. 

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Fig. 20. Southern part of distribution of the maritime striped tree squirrel 
species, Tamiops maritimns, as determined by plotting collecting localities of mate- 
rial examined. Subspecies: A, maritimus; B, monticolus; C, hainanus; and D, moi. 

Tamiops maritimus maritimus (Bonhote) 

Seiurus macclellandi maritimus Bonhote, 1900, Ann, Mag. Nat. Hist. (ser. 7), 

5, p. 51. 
Seiurus mjxcclellandi formosanus Bonhote, 1900, Ann. Mag. Nat. Hist. (ser. 7), 

5, p. 52. 
Tamiops sauteri J. A. Allen, 1911, Bull. Amer. Mus. Nat. Hist., 30, p. 339. 


Types. — Sciurus m. maritimus, BM No., adult from 
Foochow, Fukien, China, collected April 1893 by C. B. Rickett; 
formosanus, BM No., adult female from northern For- 
mosa, collected April 1862 by Robert Swinhoe; sauteri, AMNH No. 
31621, adult male from Chip-Chip, northern Formosa, collected 
November 1908 by H. Sauter. 

Material examined, from F^ikien Province, China. — Yuki (AMNH), 
three, (CNHM), two, (MCZ), two; Fuching Hsien (AMNH), 10, 
(MCZ), one; Kucheng (CNHM), one; 70 miles S.W. of Yen-Ping- 
Fu [Yenping], 500 feet (USNM), six; "N.W. Fukien" (MCZ), two; 
"Fukien" (CNHM), two. 

Matrial examined, from Formosa. — Kagi Dist., Central Formosa 
(CNHM), one; Chip-Chip (AMNH), five; Shineigun (MCZ), one; 
"Formosa" (CNHM), one. 

G. M. Allen (1940) reports a specimen in the British Museum 
from Chiong Lok, 100 miles west of Swatow, Kwangtung, and we 
have included this on our map. 

Original description. — "This is the form which most nearly ap- 
proaches the typical Sc. MacClellandi, from which it differs in being 
far greyer and more concolorous. The median dorsal stripe is by no 
means well marked and very short, not being continued to the root 
of the tail. The two subdorsal [inner light] stripes are of the same 
colour as the back, while the outermost light stripes are of a dull 
white, very narrow and short, not being continued to the root of the 
tail, and only starting at the shoulder." 

Tamiops m. maritimus, as we know it here, appears to occupy 
the vegetation area which Chi-Wu Wang (1956) has mapped as 
Rain Forest. 

Tamiops maritimus monticolus (Bonhote) 

Sciurus macclellandi monticolus Bonhote, 1900, Ann. Mag. Nat. Hist. (ser. 7), 
5, p. 52. 

Type.— BM No., adult from Ching Feng Ling, 1500 to 
2000 feet, 100 miles northwest of Foochow, Fukien, China, collected 
December 1896 by C. B. Rickett. 

Material examined, from Fukien Province, China. — Chungan 
Hsien (AMNH), 17, (MCZ), four; mountains near Yenping (AMNH), 
11, (MCZ), one, (USNM), four, (CNHM), six. 

Material examined, from Kwangsi. — Yao-shan (CNHM), two. 

Because of loss of data from our own examinations of material in 
European museums on this species, we are departing in this instance 


from the practice of citing and mapping localities only of material 
examined, and are including here and mapping the additional records 
reported by G. M. Allen (1940, pt. 2, p. 684): 

Material examined, from Fukien. — Pucheng (BM), two; Kuaton 
[Kaotien] (BM), eight; Chungfengling [Chingfengling] (BM), eight; 
Kienyang (BM), one; Tingchow (BM), two. 

Material examined, from Anhwei. — Chinteh [Tsingteh] (BM), 10. 

Material examined, from Hupeh.^ — Changyang (BM), one. 

Material examined, from Chekiang. — Ningpo (BM), one. 

Original description. — "Brighter than [maritimus] to which it is 
most nearly allied, and from which it differs in having the median 
dorsal stripe more distinct and always continued to the root of the 
tail. The outermost light stripes are very broad and distinct and 
continued to the root of the tail." 

Discussion.— We find contrary to G. M. Allen (1940, pt. 2, p. 684) 
that Bonhote was right in differentiating between the coastal samples 
of this species and those inland in the mountains. However, he did 
err in trying to distinguish them primarily upon a character of the 
middorsal stripe. The better character which he merely mentions 
is that the outer pair of dorsal light stripes continues posteriorly to 
the base of the tail and are broader and more distinct throughout 
than in maritimus. Bearing in mind, of course, the pelage differences 
related to season, we note that in the summer monticolus material 
from the mountains about Chunganhsien, the dark stripes are gen- 
erally blacker than in the winter material from there; although there 
is also one taken on December 9 from the mountains near Yenping, 
on which the dark stripes are quite as black as any with summer 
dates. Comparing only winter series, six January and three March 
maritimus specimens from Fuchinghsien with five December, one 
January, two February, and one March monticolus ones from near 
Yenping, we find the two series easily distinguished from each other. 
Eight of the nine Yenping ones in winter pelage are individually dis- 
tinguished when mingled in a series of maritimus. The ten maritimus 
from Fuchinghsien and three from Yuki in winter pelage remain 
equally distinguishable when placed in the series of eight Yenping 
monticolus. The Yenping material in winter pelage is also consist- 
ently redder in the dark stripes and its inside light stripes are lighter 
and huffier than the nape whereas those of the Fuchinghsien material 
are the olive gray of the nape. These latter characters are not dis- 
played by the four Chunganhsien winter pelage examples of mon- 
ticolus, however. This may be a formerly more distinct northern 


subspecies of maritimus in the process of being obliterated, or our 
material may be from a zone of intergradation between the two good 

Tamiops maritimus monticolus appears to occur in an area which 
coincides rather well with the extent in eastern China of two of Chi- 
Wu Wang's (1956) general vegetation types: 1. Evergreen broad- 
leaved forest of evergreen oaks, schima and laurels, with Pinus mas- 
soniana in secondary stands. 2. Mixed mesophytic forest. 

Dimensions. — In six maritimus from 500 feet elevation 70 miles 
southwest of Yenping, Fukien, taken in November and December of 
1921, and four monticolus taken in April of 1922 at 2000 feet eleva- 
tion near Yenping, all by Arthur de Carle Sowerby, hind foot and 
head-and-body size show no differentiation, but five of the maritimus 
have tails recorded between 120 and 125 mm., whereas the longest 
three of the monticolus range from 110 to 117 mm. 

Tamiops maritimus hainanus (J. A. Allen) 

Tamiops macclellandi hainanus J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 

22, p. 476. 
Tamiops macclellandi riudoni J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 

22, p. 477. 
Tamiops macclellandi laotum Robinson and Kloss, 1922, Ann. Mag. Nat. Hist. 

(ser. 9), 9, p. 92. 

Types. — Tamiops m. hainanus, AMNH No. 26664, an adult fe- 
male from Lei Mui Mon, mountains of Hainan Island, China, col- 
lected December 31, 1902, by agents of Alan Owston; riudoni, 
AMNH No. 26672, an adult male from Riudon, lowlands of Hainan 
Island, China, collected March 5, 1903, by agents of Alan Owston; 
laotum, BM No., adult male from Pak Hin Bun, Mekong 
River, Laos, collected March 2, 1920, by Herbert C. Robinson. 

Material examined, from Hainan, China. — Nodoa (AMNH), 57, 
(CNHM), 14, (MCZ), six, (USNM), one; Nam Phong (AMNH), 
10, (CNHM), four, (USNM), one; Riudon (AMNH), two; Lei Mui 
Mon (AMNH), nine, (USNM), two; Kachek (USNM), one. 

Material examined, from Tonkin, Vietnam. — Pakha [Pa Kha] 
(CNHM), one; Muong Boum (CNHM), six; Lieng San [Leng Sang] 
(CNHM), five; Nong Lum (CNHM), three; Chapa [Cha Pa] 
(CNHM), seven, (USNM), two, (MCZ), one; Phong Tho (CNHM), 
one; Muong Mo (CNHM), seven, (USNM), one, (MCZ), one; Ba 
Nam Chi (CNHM), one; Lai Chau (CNHM), two, (USNM), two, 
(AMNH), one; Muong Moun (CNHM), five, (AMNH), two; Pa 


Ham (CNHM), two; Bac Tan Tray (AMNH), one, (ANSP), one; 
Moung Mouen (ANSP), two. 

Material examined, from Annam, Vietnam. — Hoi Xuan (CNHM), 

Material examined, from Laos. — Lao Fou Tchay (CNHM), one; 
Phong Saly (CNHM), five; Muong Yo (CNHM), five; Bonn Tai 
[Bun Tai] (CNHM), one; Lo Tiao (MCZ), two; Xieng Khouang 
(AMNH), two, (MCZ), three; Tha Ng'on, Vientiane (CNHM), one; 
Saravane (CNHM), one; Phu Kobo (MCZ), three; Col de Taloun 
(MCZ), two; "Indochina" (CNHM), one. 

The extensive series in winter pelage now available from Hainan 
and Tonkin show no color difference from the winter pelage material 
of maritimus, but as G. M. Allen (1925, p. 7) pointed out and Osgood 
(1932, p. 290) amplified, there is a size difference. Hind feet in mari- 
timus measure about 32 to 35 mm. in the dried skins, and those of 
hainanus measure about 28 to 30 mm. in the dried condition. The 
skulls of maritimus are also larger. No other examples of the reddish 
riudoni have appeared to justify the supposition that this represents 
a lowland race, and the type apparently is a strongly erythristic in- 

Robinson and Kloss (1922) attributed no diagnostic characters to 
their laotum, and although Osgood (1932, p. 291) retained it as "well 
characterized by very pale color which reaches its extreme develop- 
ment in southern Laos," he had only one specimen from southern 

Tamiops maritimus moi (Robinson and Kloss) 

Tamiops macclellandi moi Robinson and Kloss, 1922, Ann. Mag. Nat. Hist, 
(ser. 9), 9, p. 92. 

Type.— BM No., adult male from Langbian Peaks, 
5500 to 6500 feet, southern Annam, collected April 25, 1918, by 
C. Boden Kloss. 

Material examined, all from southern Vietnam. — Pie de Langbian 
(MCZ), four; Langbian Peak (MCZ), two, (CNHM), 10; Dalat 
(CNHM), one; Gougah (MCZ), one, (CNHM), two. 

Original description. — "Like T. m. laotum, but darker above; up- 
per parts more suffused with ferruginous, so that in addition to being 
more rich-coloured generally, the yellow stripes are ochraceous in- 
stead of buff, while the rump and the outer pair of dorsal stripes are 
a brighter brown." 


Discussion. — This is only a faintly distinguishable subspecies. 
The winter pelage is like that of hainanus excepting for the two 
dark paired dorsal stripes being faintly redder. We find that the 
hind foot of moi is larger than that of hainanus, however, in the seven 
MCZ specimens, which range between 31 and 32.2 mm. in the dried 
skins. The skull does not appear to be appreciably larger than that 
of hainanus. 


Genus DREMOMYS Heude 

Dremomys Heude, 1898, Mem. Hist. Nat. Emp. Chinois, 4, part 2, p. 54. 
Zetis Thomas, 1908, J. Bombay Nat. Hist. Soc, 18, p. 244. 

Type species. — Dremomys, Sciurus pernyi Milne-Edwards; Zetis, 
Sciurus rufigenis Blanford. 

Definition. — The genus Dremomys consists of four species of the 
Indochinese Subregion, lokriah, pernyi, rufigenis, and pyrrhomerus, 
and one species of the Malaysian Subregion, everetti. See Figure 21. 

Diagnosis. — The genus Dremomys possesses the following char- 
acteristics: (1) There is one bony septum crossing the chamber of the 
auditory bulla. (2) The teeth are orthodont or slightly proodont. (3) 
The length of the nasal bone exceeds the least interorbital breadth. 
(4) The upper cheek teeth are not specialized with an extremely deep 
valley that remains as a dirt-filled lake after the cusps have been 
planed away by use. (5) The coronoid process of the mandible is 
well developed and strongly falcate. (6) The baculum is composed 
of two separate parts, shaft and blade. (7) The dorsal pelage has no 
longitudinal light stripes. 

The genus Dremomys may be distinguished from other genera of 
Sciurinae of the Indian and Indochinese subregions by the above 
characteristics as follows: from Ratufa by 1, 2, 3, and 6; Funambulus 
by 5, 6, and 7; Callosciurus by 3; Tamiops by 3 and 7; Menetes by 4, 
5, and 7; and Sciurotamias by 1 and 2. 

Systematic history. — Dremomys was poorly characterized by Heude. 
Thomas (1908) discussed the genus under the name Zetis in greater 
detail, and gave a key to the species (pp. 248-249). Later Thomas 
(1916a) published another synoptic key using the generic name 
Dremomys. Pocock (1923) and Ellerman (1940) gave the genus full 
standing, and Archbold and Tate (1935, p. 1, footnote 1) designated 
Dremomys pernyi as the genotype. Besides the species pernyi, Heude 
included three other species in the genus which he described as new: 
saltitans, collaris and latro. The animals to which these three names 
were applied, however, have since been placed in the genus Sciuro- 





^^Y OF 3EN(^Al. 


■ lokriah 
^ pernyi 





Fig. 21. Composite, slightly generalized distribution of the five species of the 
long-nosed squirrel genus Dremomys. 

tamias (G. M. Allen, 1940, p. 646). It was recognized in the early 
nineteen hundreds that Dremomys was the proper genus for the spe- 
cies lokriah and rufigenis, and various authors have at one time or 
another (correctly) placed the species everetti and pyrrhomerus in this 
genus. These four species and the original one pernyi constitute the 
genus Dremomys. 

Range. — On the continent it is a wide-ranging genus, extending 
from Nepal in the Himalayas 2000 miles east across the breadth of 
China to the coastal province of Chekiang, and from only 250 miles 
above the equator in the Malay Peninsula north through Siam, 
Burma, Indochina and China nearly 2000 miles to northern Szech- 
wan. It occurs also on Borneo, Hainan, and Formosa. 

Dremomys is a long-nosed squirrel, characteristically with plain, 
drab dorsal pelage. It is generally reported to be terrestrial or semi- 
terrestrial in habit, and to occur only in the mountains or hills where 

Fig. 22. Skull and left mandible of the Himalayan long-nosed squirrel, 
Dremomys lokriah, AMNH No. 114936, Xl. 

Fig. 23. Skull and left mandible Xl of the Chinese long-nosed squirrel, 
Dremomys pernyi, AMNH No. 43999, but lower jaw of AMNH No. 114938. 


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there is some forest. Its species all have a small, rather inconspicu- 
I ous, whitish or buffy patch of contrasting pelage immediately behind 
the ear. 


1 . No yellow, orange, or red colored pelage everetti (extraterritorial) 

Some yellow, orange, or red colored pelage 2 

2. Under surface of tail not red 3 

Under surface of tail brilliantly red full length 4 

E 3. Ventral pelage of body all gray at bases but all yellow or orange tipped (No red- 

f dish brown anal pelage patch) lokriah 

I Ventral body pelage all white tipped or partly yellow tipped but color patch 

I of Burnt Sienna present about anus pernyi 

4. Sides of head (but not the top of it nor the throat) bright red, flash mark on 

hip inconspicuous rufigenus 

No red on head, flash mark on hip conspicuous pyrrhomerus pyrrhomerus 

Head entirely red, flash mark on hip conspicuous (sometimes spreading over 

entire thigh) pyrrhomerus riudonensis 

. Flash marks on cheeks and hips inconspicuous but throat red. 

pyrrhomerus gularis 

Dremomys lokriah (Hodgson) 

Definition. — The species Dremomys lokriah includes the subspecies 
lokriah, macmiUani, garonum, and pagus, and the named forms in- 
cluded in these as synonyms. The species distribution is mapped in 
Figure 24. 

Diagnosis. — The species lokriah is characterized by (1) The dorsal 
pelage is plain, dull agouti without red on cheeks or hips. (2) The 
ventral pelage is yellow or orange without any reddish-brown perineal 
patch. (3) The under side of the tail is not red. (4) There are white 
tips to the tail hairs. 

The species lokriah is distinguished from the other species of its 
genus by the above characters as follows: from pernyi by 2; from 
rufigenis by 1, 2, and 3; from pyrrhomerus by 1, 2, and 3; from everetti 
by 2 and 4. 

Relationships to other species. — Zahn (1942) includes in the species 
lokriah approximately the geographic races that we assign to it here, 
but he adds to it a lumping of the races which previously belonged 
to a separate species, Dremomys pernyi. Anthony (1941, p. 91) took 
samples of both lokriah and pernyi in the vicinity of Imaw Bum in 
northeast Upper Burma, and they differ sharply at this locality. 
The ventral pelage of the pernyi material is white, and that of the 
lokriah Orange Ochraceous. The skulls of pernyi are trenchantly 



long-snouted; those of lokriah are relatively short. We have also 
seen material representing both species, from 6000 feet in the Adung 
Valley about 130 miles farther north ; and here again both pelage and 
skull showed the same trenchant differences. The total amount of 
this material seen is, of course, small (three lokriah, four pernyi), and 
the localities constitute the known eastern extremities of the range 
of lokriah. At the western limit of the range of pernyi some 300 
miles within the range of lokriah, there does appear to be some in- 
teresting similarities between the local races of the two species. See 
the account of D. I. macmillani for details. 

Dremomys lokriah lokriah (Hodgson) 

Sciurus lokriah Hodgson, 1836, Jour. Asiatic Sec. Bengal, 5, p. 232. 
Dremomys lokriah bhotia Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, 

p. 426. 
Dremomys lokriah subflamventris Thomas, 1922, Jour. Bombay Nat. Hist. Soc, 

28, p. 429. 

Fig. 24. Geographical distribution of the Himalayan long-nosed squirrel, 
Dremomys lokriah, as shown by collecting locality records of material examined. 
Dotted lines enclose mountainous areas probably inhabited by this species, but 
should not imply that it crosses the Salween River. Subspecies: A, lokriah; 
B, macmillani; C, garonum; and D, pagus. 


Types. — Sciurus lokriah, BM Nos. and old male 43.1. 
12.56 (cotypes) both from Nepal; bhotia, BM No., old 
male collected at Sedonchen at 6500 feet elevation, Sikkim, India, 
November 14, 1914, by C. A. Crump; subflaviventris, BM No. 79.11. 
21.351, old adult from Assam, collected by John McClelland. 

Material examined. — "Nepal" (BM), seven, (RNH), one; "Tibet" 
(RNH), one; Chandragiri Pass, 7000 feet, Nepal (BM), one, (USNM), 
four; Hathiben, Nepal (BM), one; Godaveri, 7000 feet, Nepal 
(USNM), one; Sattha, 2 miles northwest of Gorkha, Nepal (BM), 
one, (CNHM), one; "Sikkim," 9000 feet (BM), two; Sukiapokhri 
[Sookia Pokhari], 7000 feet, Darjeeling (BM), three; Darjeeling, 
7000 feet, Sikkim (BM), two; "Jeluk," 9800 feet, Sikkim (USNM), 
one, (CNHM), 10; Gopaldhara, Rungbong Valley, Sikkim (CNHM), 
one; Lachen, 8800 feet, Sikkim (CNHM), one, at 6000 feet (BM), 
one; Gangtok, 7000 feet, Sikkim (BM), one; Ringin, 6000 feet 
Sikkim (BM), one; Chungtang, 5350 feet, Sikkim (USNM), one, 
(CNHM), six; Sedonchen, 5500 to 6500 feet, Sikkim (BM), three; 
Lachung, 8000 feet, Sikkim (CNHM), one; Dreyi, 5140 feet, Mishmi 
Hills, Tibet (BM), seven; Nyetmaw River, 8600 feet, Burma 
(AMNH), one; Mt. Imaw Bum, 9000 feet, Burma (AMNH), one; 
Taron Valley, 7000 feet (BM), one; Akhe Triangle, 3500 feet (BM), 
one; Adung Valley, 6000 feet (BM), two, (CNHM), one. 

Original description. — "Above, saturate brown, tipped with in- 
tense orange; below, and the thighs, deep orange. Tail concolorous 
with body above, distichous, flattened, and broad, with a double 
margin of black and hoary." 

Type descriptions. — The types of lokriah, like subflaviventris, bho- 
tia, and garonum have the underparts dull mustard yellow, in con- 
trast to the yellowish white of macmillani. The Leiden (RNH) 
specimens are dark gray-brown, a grizzle of buff tips in gray bases, 
the deep bases smoky. The underparts deep orange. No orange at 
base of tail. Throat buffy white. Dorsal color unchanged on hands 
and feet. 

Pelage color. — The color of the ventral pelage being diagnostic, it 
seems worth while to mention that that of the United States National 
Museum material from Nepal was recorded as Ochraceous Orange. 
That of the Chicago Natural History Museum material from Sikkim 
was with very little variation, Salmon Orange, excepting in one or 
two of the adults where it approached Xanthine Orange closely. 
That of the American Museum material from northern Burma is 


Ochraceous Orange. Salmon Orange is, of course, very near Ochra- 
ceous Orange. 

Discussion. — The Chicago Natural History Museum series of 
D. lokriah from Sikkim convinced us that there are not two good 
geographic races involved there, separated by the Teesta River as 
proposed by Wroughton (1916c). In the attenuated distribution of 
the race D. I. lokriah across the face of the Himalayas, the Dafla, 
Abor, and Mishmi hills, and into the mountain massif of northern 
Burma, one would look for variation. The American Museum ma- 
terial available from very northern Burma seems to provide a re- 
duction in size, but the sample (2) is much too small to demonstrate 
geographic subspeciation. 

Material from the range of D. I. lokriah as we recognize it here, 
is rather scarce, and it samples widely separate localities. Neverthe- 
less, the pelage characters of the Adung Valley, Burma, specimen, 
for example, make it inseparable from the series of skins from Sik- 
kim. In skull characters we find virtually no geographic variation 
between the subspecies of lokriah, but in the very small samples of 
skulls that we have measured from the eastern and western extrem- 
ities of the range of D. I. lokriah, the auditory bullae run proportion- 
ally smaller than in Sikkim. The sample from the western extremity 
(Nepal) also has a proportionally longer orbit, and one wonders if 
the Arun Valley is not a rather effective barrier to any east-west 
passage of this squirrel, isolating the main Nepalese population from 
interbreeding with the population east of this river. 

Habits. — Lord Cranbrook {in Kinnear, 1934) noted that Dre- 
momys pernyi and D. [lokriah] seemed in the Adung Valley to replace 
Callosciurus erythraeus where the forest changed from subtropical to 
a more temperate type with conifers. This is quoted in the present 
paper in the account of C. flavimanus quinquestriatus. 

Another observer who collected this species comments: "Common 
in all the forests from 5,000 to 9,000 feet. Lives in holes in trees, 
generally low down and is frequently seen on the ground, feeding on 
fallen nuts and berries. As a rule it is silent but on occasion utters 
a loud cackling note. When approached it hides itself by lying 
flat along a branch, and does not attempt to leave the tree unless 
really frightened." (C. A. Crump in Wroughton, 1916a, p. 488) 

Dremomys lokriah garonum Thomas 

Dremomys lokriah garonum Thomas, 1922, Jour. Bombay Nat. Hist. Soc, 28, 
p. 430. 


Type.— BM No., adult male from Tura, 1200 feet, Garo 
Hills, Assam, collected February 25, 1920, by H. W. Wells. 

Material examined, all from Assam, India. — Tura Mt., Garo Hills 
(CNHM), one; Duragiri [Darugiri] 3000 feet, Garo Hills (BM), one; 
"Rajapara," 600 feet, south Kamrup (BM), one; Umran, Khasi Hills 
(AMNH), two; Bara Pani [Borpani], Khasi Hills (AMNH), two; 
Nongpoh, Khasi Hills (AMNH), three, (CNHM), one; Pynursla, 
Khasi Hills (CNHM), one; Mawphlang, Khasi Hills (CNHM), 34; 
Mawlyngkueng, Khasi Hills (CNHM), four; Laitlynkot, Khasi Hills 
(CNHM), one; Cherrapunji, Khasi Hills (CNHM), two; and "Kon- 
shnong," 3000 feet, Jaintia Hills (BM), three. 

Original description. — "Size about as in . . . lokriah, or slightly 
smaller. Colour above as in D. I. [lokriah] but below ... it is far paler 
and more yellow, nearly matching Ridgeway's 'orange-buff.' Buffy 
of underside narrowed. . . . Readily distinguishable by the paleness 
of its lower surface." 

Dremomys lokriah pagus Moore 

Dremomys lokriah pagus Moore, 1956, American Mus. Nov. No. 1816, p. 1. 

Type.— AMNH No. 163479, an adult male from 1400 meters ele- 
vation on Mt. Victoria in the Pakokku Chin Hills of western Burma, 
collected March 23, 1938, by Gerd Heinrich. 

Material examined. — Mt. Victoria, Pakokku Chin Hills, Upper 
Burma (AMNH), 18; Sangau, Lushai Hills, Assam (CNHM), six. 

Pelage coior.^Dorsal pelage lifte that of subspecies lokriah and 
garonum; ventral pelage with bases of Slate Gray and tips of Ochra- 
ceous-Buff . Colored hair tips are longer on the throat so that Slate 
Gray does not show through there; in the rest of the ventral pelage 
it does. 

Diagnosis. — Dremomys I. pagus differs from subspecies lokriah 
and garonum by paler, more yellow ventral pelage; from macmillani 
by lacking a distinct anal area pelage color patch, and usually by 
lacking a middorsal blackish stripe. 

Discussion. — This is the southernmost geographic race of D. lok- 
riah and its type locality is our southernmost locality record for the 
species. It is only in contact with macmillani and presumably inter- 
grades with it in the Chin Hills and to some extent the Lushai Hills — 
a third of the Lushai Hills specimens having the blackish middorsal 


Dremomys lokriah macmillani (Thomas and Wroughton) 

Dremomys macmillani Thomas and Wroughton, 1916, Jour. Bombay Nat. 
Hist. Soc, 24, p. 238. 

Type.— BM No., young male from Tatkon, 250 feet, 
west bank of the Chindwin River, opposite Kindat, Burma, collected 
June 27, 1914, by S. A. Macmillan. 

Material examined, all from Assam, India .^ — Mokokchung, 4000 
feet, Naga Hills (BM), one; "Naga Hills," 5000 feet (BM), two; 
Takubama, Naga Hills (CNHM), five; Karong, Manipur (CNHM), 
12; five miles north of Imphal, Manipur (USNM), one; and Chin 
Hills, 5000 feet, 50 miles west of Kindat, Burma (BM), three. 

Original description. — "Size about as in . , . lokriah . . . colour 
above dark coarsely grizzled olive-grey, clearer on fore-back, suffused 
with dull tawny on crown, nape, and hind-back; a distinct narrow 
black median line present . . . from the back of the nape to the loins. 
. . . Undersurface bright buffy, lighter anteriorly and on inner side 
of the forelimbs, darkening to cinnamon buff on the inner side of the 
hind limbs. . . . Anal region and base of tail beneath rich ochraceous- 
rufous. . . . Postauricular patches prominent, deep ochraceous buffy. 
. . . Tail hairs ringed with black and pale buffy, . . . tips white; no 

rufous . . . along underside distinguishable [from lokriah] 

by its distinct dorsal black line, its greyer general colour, the more 
completely buffy hairs of the underside, and by the greater promi- 
nence and buffy colour of the postauricular patches." 

Discussion. — Takubama is evidently in the area of intergradation 
between D. I. lokriah and D. I. macmillani. Only one of the five from 
there possessed the middorsal stripe, and two the ventral pelage of 
macmillani; whereas two have pelage identical to that of the D. I. 
lokriah material from Sikkim, and one has a venter like the D. I. lok- 
riah material from Imaw Bum and Nyetmaw River. From the ma- 
terial examined, Kohima, Karong, and Imphal are, however, firmly 
within the range of macmillani. 

Examples of Dremomys pernyi have also been examined from 
Takubama, Karong, and Kohima, and there are the following simi- 
larities between the two local subspecies of the two species. Of the 
dozen specimens of macmillani from Karong, three have pure white 
pelage streaks occurring both in the middle of the throat and the 
middle of the chest in each individual. Half of the dozen Karong 
specimens have their entire ventral pelage as pale as Light Ochra- 
ceous-Buff (the others: 5 Ochraceous-Buff, 1 Orange-Rufous). The 
skin of one adult female D. pernyi howelli in the Takubama material 


(CNHM 76344) is in its ventral color, intermediate between the spe- 
cies lokriah and pernyi. In the ratio of the orbito-nasal length to 
the greatest length of the skull, however, and in general character 
of the skull, this specimen is well outside of the range of the species 
lokriah and in that of the longer-nosed pernyi. The local race of the 
white-bellied species, Dremomys pernyi howelli, which inhabits the 
same area as Dremomys lokriah macmillani, like the latter is distin- 
guished from its own conspecific relatives in part by a blackish mid- 
dorsal stripe. Dremomys lokriah macmillani is the only geographic 
subspecies of lokriah which possesses a characteristic color patch 
about the anus, which does, however, characterize the entire species 
pernyi. One might suspect this variable subspecies D. I. macmillani 
which so resembles D. pernyi howelli, of being a product of inter- 
gradation between the two species. Study of the skull material rep- 
resenting macmillani reveals no real suggestion, however, of any such 
intergradation or of species cross. There is a typical lokriah lokriah 
skin wearing the same number (CNHM 76312) as a typical pernyi 
howelli skull from Takubama in the Naga Hills, but this is assumed 
to be mixing of collected specimens. 

Dremomys pernyi (Milne-Edwards) 

Definition. — The species pernyi is constituted by subspecies pernyi 
howelli, flavior, senex, owstoni, and calidior, and see the synonyms 
listed under these. The distribution of this species is mapped in 
Figure 25. 

Diagnosis. — Characteristics of the species pernyi are: (1) The dor- 
sal pelage is plain agouti gray without flash mark on hip or red cheek. 
(2) The ventral pelage is marked by a reddish brown patch about the 
anus. (3) The ventral pelage is whitish on thorax and abdomen. 
(4) The ventral pelage of the tail is a faintly buffy gray. (5) The 
tail hairs are tipped with white. 

The species pernyi is distinguished from other species of Dre- 
momys by the above characters as follows: from lokriah by 2, 3, and 
4; rufigenus by 1, 2, and 4; pyrrhomerus by 1, 2, and 4; and everetti 
by 1 and 5. 

Relationships to other species. — As may be seen in comparing Fig- 
ures 24 and 25, this species replaces the species lokriah on the east, 
and is sympatric with it in northern Assam and Burma. In central 
and eastern China there are so few collections that it is difficult to 
assess its relationships with species pyrrhomerus, with which it does 
in some manner appear to share this vast area. Both were collected 



at Ichang, Hupeh; Pien Ngai, Szechwan; and Suiyang, Kweichow. 
To the south pernyi is replaced by species rufigenis. The ranges of 
three species, lokriah, pernyi, and rufigenis seem to overlap broadly 
in upper Burma. 

Our study of this species with the rusty anal patch, reveals six 
geographic races as listed above. In this we find ourselves in fairly 
close agreement with Allen (1940, p. 646), but differing considerably 
with Zahn (1942, pp. 124, 127, 182), who recognizes only pernyi and 
senex, and Ellerman and Morrison-Scott (1951, p. 492), who admit 
only pernyi, imus, and owstoni. 

Dremomys pernyi pernyi (Milne-Edwards) 

Sciurus pernyi Milne-Edwards, 1867, Rev. et Mag. Zool. (ser. 2), 19, pi. 19. 
Dremomys pernyi griselda Thomas, 1916, Ann. Mag. Nat. Hist. (ser. 8), 17, 
p. 392. 

Dremomys pernyi lichiensis Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, 

p. 403. 
Dremomys rufigenis lentus A. B. Howell, 1927, Jour. Washington Acad. Sci., 

17, p. 80. 

Types.— Sciurus pernyi MNHN No. 1868-1331 (200), adult from 
"les montagnes de la principaut^ de Moupin," Szechwan, China, col- 
lected by Perny; griselda, BM No., young female from 
Nagchuka [Hokow], 10,000 feet, Szechwan [Sikang], China, collected 
May 25, 1911, by F. M. Bailey; lichiensis, BM No., young 
male from LiChiang Range, latitude 27° 20' N., Yunnan, China, ele- 
vation 10,000 to 12,000 feet, collected July, 1918 by G. Forrest; 
rufigenis lentus, USNM No. 240384, adult male from Wen Chuan 
Hsien, 6000 feet, Szechwan, collected August 14, 1924, by D. C. 

Material examined, from Sikang. — Shoo-o-lo [Siolo or Singolo] 
West Szechwan (BM), one; Ramala Pass [Lamaya], Szechwan 
(USNM), one; Ta Chien Lu [Tatsienlu], 12,000 feet (CNHM),one, 
(MCZ), one; Mi-li (CNHM), four; Baurang (CNHM), one; Ti-yu 
Gomba [Dayul Gomba], 12,950 feet (CNHM), one; Yulong-Vilang 
[Yulong] (CNHM), one; Mupin (USNM), two; "Yao Gi" near Mu- 
pin (USNM), one; Tang Gu [Tangshu], near Gieu Long Shien, Tibet 
[Sikang] (USNM), four; "Wa Hu Pass," 14,000 feet, Tibet (USNM), 

Fig. 25. The species range of the Chinese long-nosed squirrel, Dremomys 
pernyi, as shown by plotted collecting localities of material examined. Open dots 
in this figure represent material intermediate in character between subspecies. 
Subspecies: A, pernyi; B, howelli; C, flavior; D, senex; E, calidior; and F, owstoni. 


one; Nagchuka [Hokow] (BM), one; "Yao Chao," east of Mekong 
River, 13,000 feet (BM), one. 

Material examined, intermediate with D. p. howelli, from Si- 
kang.— Mekong-Salwin Divide, latitude 28° 20' N., 9000-10,000 feet 
(BM), seven. 

Material examined, from Szechwan. — 25 miles west of Wench- 
wan, 7000 feet (AMNH), four; "Wu-chi" (CNHM), four; "Mei- 
peng" (CNHM), one; "Szechwan" (USNM), one. 

Material examined, intermediate with D. p. flavior, all from Yun- 
nan.— Nguluko (USNM), one; Lichiang [Likiang], 8000-12,000 feet 
(CNHM), eight, (MCZ), two, (USNM), seven, (AMNH), four; 
Lichiang Range, latitude 27° 30' N., 9000-11,000 feet (BM), 19; 
"Teshweko," 11,000 feet, Likiang Range (USNM), one; "Lhotan," 
12,000 feet. Western Likiang Snow Range (USNM), one; Northern 
slope Likiang Snow Range (AMNH), one. 

Material examined, from Yunnan.^ — "Hofuping Mts.," Kekong 
Valley (USNM), three; Li-tien and Wieshi Pass (AMNH), two; 
"Mts. of Yangtze, Mekong Valley" (USNM), one; Mekong- Yangtze 
Divide, 7000 to 8000 feet, latitude 27° 30' N. (BM), one; Mekong 
Valley, 7000 feet, at latitude 28° N. (BM), two; Mekong-Salween 
Divide, 7000 to 10,000 feet, latitude 28° 20' N. (BM), two; Tse-kow 
[Tseku], N.W. Yunnan (BM), one; Yung-ning (CNHM), four. 

Type description. — Milne-Edwards {loc. cit.) published only the 
colored plate of the type of pernyi with no description or other de- 
tails in words. The present (1951) appearance of the type is dull gray 
brown, a grizzle or mixture of yellowish and blackish brown. This 
color is uniform all over the back, limbs and feet, and onto the head. 
The tail is only slightly darker, due to a greater amount of black. 
Claws of the forepaws are rather long. The underparts are white 
from chin to vent. The anal region and inner sides of the hind limbs 
posteriorly are bright rusty brown. 

Discussion. — This is a fairly distinct, large, northern race, dis- 
tinguished from its conspecific neighbors on the south and west by 
its lighter gray dorsal pelage and tail. 

The abundant material from the high mountain peninsula formed 
by a long loop of the Yangtze about Likiang has caused much con- 
fusion. Most of this material we find indistinguishable from pernyi 
of Sikang to the north, but some of it seems intermediate between 
pernyi and flavior; and it seems reasonable on geographic grounds 
to regard material of the species D. pernyi in the Likiang area as 


Just north of Likiang the narrow area between the Yangtze and 
Salween rivers, divided by the Mekong, is an area of intergradation 
between D. p. pernyi and D. p. howelli, whence the material is also 
abundant and may appear to represent either race. Thomas (1922, 
p. 400) came to regard this small area as that of typical D. p. pernyi, 
with imus across the Mekong to the west and griselda across the 
Szechwan (Sikang) border to the east, and with griselda 30' of lati- 
tude (34.4 miles) to the south and griselda 20' of latitude (23 miles) 
to the north. Whether the type specimen came from that area or 
more probably from near Muping, Sikang, the area occupied by a 
recognizable geographic subspecies which can logically bear the name 
D. p. pernyi is to the north and east in Sikang and Szechwan, and 
the characters which distinguish it are those which Thomas applied 
to his D. p. griselda. 

Dremomys pernyi howelli Thomas 

Dremomys pernyi howelli Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, 

p. 401. 
Dremomys pernyi m^ntosus Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, 

p. 401. 
Dremomys pernyi imus Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 402. 

Types. — Dremomys p. howelli, BM No., old male from 
Ma-Chang-Kai, 6500 feet, about 25 miles S.W. of Tengyueh, upper 
Shweli River, extreme western Yunnan, China, collected June 4, 
1912, by E. B. Howell; mentosus, BM No., old female from 
5000 feet in the Chin Hills 65 miles [6 in type description] southwest 
of Kindat, Upper Burma, collected May 13, 1915, by J. M. D. Mac- 
kenzie; imus, BM No., old male from 7000 feet on the west 
slope of Mt. Imaw Bum, Upper Burma, collected October 21, 1919, 
by F. Kingdon Ward. 

Material examined, from Yunnan. — Ma-Chang-Kai, 6500 feet, 
Tengyueh (BM), six; Shweli-Salween Divide, 7000-10,000 feet (BM), 
two; Tai-Ping-Pu, 7000 feet, Schweli River (AMNH), two. 

Material examined, from Upper Burma. — Taron Valley, 4500 feet 
(BM), one; Adung Valley, 6000 feet (CNHM), two; Nam Tamai 
Valley, 3000 feet (BM), two; Gangfang, 5200 feet (AMNH), one; 
Tsonma, 8300 feet (AMNH), one. 

Material examined, from Assam, India. — Manipur (BM), one; 
Karong, Manipur (CNHM), three; Kohima, Naga Hills (CNHM), 
one; Takubama, Naga Hills (CNHM), four. 


Original description. — "Colour throughout Hke . . . true pernyi, 
or very slightly more yellowish olivaceous, but fore-back in every 
specimen there is an almost imperceptible blackish dorsal line from 
one to two inches in length. Under surface as in pernyi, . . . front 
aspect of lower legs dull whitish or more or less washed with reddish. 
Tail as in pernyi." 

Discussion. — The above description is confusing, for Thomas had 
come to regard as typical pernyi certain material from what we must 
now regard as a zone of intergradation between typical pernyi and 
howelli, and also certain dark variant individuals from within the 
range of what Allen (1940, p. 648) and we recognize as subspecies 
pernyi. The characters of this proper geographic subspecies pernyi 
are in fact the ones by which Thomas (1916b, p. 392) distinguished 
his subspecies griselda. 

While the small amounts of material available from the three ex- 
tremes of the V-shaped geographic range of this subspecies can be 
to some extent distinguished and thus do provide some excuse for 
the three names, it seems quite certain that the difference will dimin- 
ish as more material even from the extremes, becomes available. 
And it should be recognized that the materials from these three ex- 
tremes of the range are more like each other than they are like the 
adjacent geographic races, pernyi and flavior. 

Dremomys p. howelli as recognized here is characterized by a 
blackish middorsal line in the pelage, fainter in the eastern leg of 
its range than in the western, but present throughout. It may be 
otherwise distinguished from both D. p. pernyi and D. p. flavior by 
generally darker dorsal pelage and tail, and by the darker, richer 
colored anal patch. In series, furthermore, the venter of flavior is 
notably whiter. 

Habits. — Lord Cranbrook (in Kinnear, 1934) noted that Dre- 
momys pernyi and D. [lokriah] seemed in the Adung Valley to replace 
Callosciurus erythraeus where the forest changed from subtropical to 
a more temperate type with conifers. This is quoted in the present 
paper in the account of C. flavimanus quinquestriatus. 

In a letter of June 10, 1961 to one of us. Lord Cranbrook contrib- 
uted the following observations from his field notes made while col- 
lecting in the Adung Valley: "No. 237. Shot on the ground in jungle. 
No. 213. Snared by native on the ground; pregnant, three young. 
January, 5000 feet." Sizes of broods of tropical squirrels are so little 
known that no records were reported for Dremomys in a recent litera- 


ture survey (Moore, 1961, pp. 21-26), hence this observation of brood 
size is of some importance. 

Dremomys pernyi flavior G. M. Allen 

Dremomys pernyi flavor G. M. Allen, 1912, Proc. Biol. Soc. Washington, 25, 
p. 178. 

Type. — MCZ, no. 13,691, young male, from Mongtz [Mengtsz], 
southeastern Yunnan, collected in 1911 by H. Orii. 

Material examined, from Yunnan. — "Yunnan" (BM), five; 15 
miles S.W. of Kunming [Yunnanfu] (USNM), two; Lung Kai [Lung- 
kai], Wuting Hsien [Wutingchow] (AMNH), three; Kao Chiao 
(AMNH), two; Feng Yang (AMNH), three; Hsin Kai [Sinkai] 
(AMNH), one; Cuchi [Kuchai] (USNM), one; Yang Wu Pa (AMNH), 
two; "Udati" near Mongtze [Mengtsz] (BM), one; Huang Jia Keo 
[Huangchiakou] (USNM), two; Tseo Jia Keo, Yunnan border south 
of Suifu (USNM), four; "Szechwan" (USNM), one. 

Material examined, from Kweichow. — Hwang Tsao Pa [Hwang- 
tsaopa] (USNM), eight. 

Original description. — "Similar to D. pernyi but smaller and yel- 
low in general coloration. The median area of the under side of the 
tail is yellowish or buffy instead of whitish. 

"Entire upper surface of the head . . . neck, body, limbs, and base 
of tail a nearly uniform grizzled buffy and black. , . . Chin, throat, 
belly and inner sides of the legs white, washed with pale buff on the 
throat. The white hairs, except on the chin, have dark slaty bases. 
Anal region pale ochraceous-rufous . . . extending onto the base of 
the tail below and the inner sides of the tibial margin of the legs. 

"... Below the central area of the tail ... is cream buff bordered 
by black and fringed with white. . . . 

"Skull. — Compared with . . . pernyi from Szechwan [Sikang] . . . 
the new race is decidedly smaller. ..." 

Discussion. — This is a quite good geographic race of D. pernyi. 
One may distinguish it from the typical D. p. pernyi to the north by 
its consistently smaller size and darker, more olive dorsal pelage. 
Agreeing with Allen (1940, p. 649) regarding the distinctness of this 
race, we differ with him, however, in that we recognize this sub- 
species only south of the Yangtze River in Yunnan and Kweichow. 
The material from Litien and Weisi Pass, which he included in this, 
we find to be intermediate between typical Dremomys pernyi pernyi 
and D. p. howelli. 


Dremomys pernyi senex (G. M. Allen) 

Dremomys senex G. M. Allen, 1912, Mem. Mus. Comp. Zool., 40, p. 229. 

Dremomys pernyi modestus Thomas, 1916, Ann. Mag. Nat. Hist. (ser. 8), 17, 
p. 393. 

Types. — Dremomys senex, MCZ No. 7582, adult female from Nan- 
tow (or Nantu), Ichang Hsien, Hupeh, China, collected February 5, 
1909 by Walter R. Zappey; modestus, BM No., adult male 
from Sui-yang, Kweichow, China, collected in April, 1898, by F. W. 


Material examined. — Ichang, Hupeh, China (BM), five, (MCZ), 
two; Suiyang, Kweichow (BM), two; Tungtze, Kweichow (CNHM), 
one; "Ta-tong Hsien," Kweichow (CNHM), one. 

Original description.— "Nearest to D. pernyi, from which it dif- 
fers in its greater size, with notably longer tail and larger skull, in 
having the postauricular patch white instead of deep ochraceous buff, 
and the median area of the ventral surface of the tail nearly uniform 
clay color instead of whitish." 

Pelage color. — "Dorsal coloring essentially as in typical D. p. 
pernyi and D. p. flavior ... a small postauricular patch of buffy or 
white. ... A narrow buffy eye-ring is present and the cheeks are 
slightly tinged with ochraceous. Throat white to roots of the hairs, 
anteriorly, but the lower throat and the belly and the forearms are 
gray-based ... a faint wash of yellowish buff . . . conspicuous on the 
border of the thighs. Tail above showing three black bands on the 
individual hairs, alternating with dull ochraceous buff to drab, and 
tipped with white; below, the white tips form an external fringe, sue- 1 
ceeded by a black border, while the central area is drabby ochraceous. 
The usual ferruginous patch is present over the anal region extend- 
ing to the upper part of the hind legs." (G. M. Allen, 1940, p. 651) 

Discussion. — Since Thomas in describing modestus commented 
that it is nearer senex, we regard the name a synonym of senex, 
although the material from Suiyang, Tungtze, and Ta-tong Hsien 
seems intermediate between senex and flavior. Obviously there is 
much yet to be learned about the distribution of senex. 

Dremomys pernyi calidior Thomas 

Dremomys pernyi calidior Thomas, 1916, Ann. Mag. Nat. Hist. (ser. 8), 17, 

p. 394. 
Dremomys pernyi chintalis Thomas, 1916, Ann. Mag. Nat. Hist. (ser. 8), 17, 1 

p. 394. 


Types. — Dremomys p. calidior, BM No., young adult 
male from Kuatun [Kaotien], northwest Fukien, China, collected by 
F. W. Styan; chintalis, BM No., young female from Chin- 
teh [Tsingteh], An-hwei, China, collected October 29, 1896, by 
F. W. Styan. 

Material examined. — Kwatum [Kaotien], Fukien (USNM), two; 
Kuatun [Kaotien], northwest Fukien (BM), 13, (MNHN), one; 
Chungan Hsien, north Fukien (AMNH), 18; "N. W. Fokien" 
(AMNH), one; Chinteh [Tsingteh], Anhwei (BM), three. 

Original description. — "General characters very much as in D. p. 
senex, but upper color a much warmer brown, approaching 'olive- 
brown.' Ear-patches mixed white and ochraceous, the bases of the 
hairs white and their tips ochraceous. Under surface whitish, but 
ordinarily with well-marked buffy thigh-patches." 

In selecting one of these two synonyms for the name of this race, 
it should have been better housekeeping to have chosen the prior 
chintalis since it actually appears first on the page. As Allen (1940, 
p. 652) strongly suggests, further collecting may reveal the necessity 
of synonymizing both under the name senex, making a single brown- 
ish subspecies in eastern China. 

Dremomys pernyi owstoni (Thomas) 

Zetis owstoni Thomas, 1908, Jour. Bombay Nat. Hist. Sec, 18, p. 248. 

Type. — BM No., adult female from Mt. Arizan, central 
Formosa, collected December 6, 1906, by A. Owston. 

Material examined. — Mt. Arizan, Central Formosa (BM), seven; 
"Central Formosa" (CNHM), three. 

Original description. — "Most nearly allied to lokriah, but larger 
and darker colored. . . . Under surface broadly and conspicuously 
washed with yellow or orange, the bases of the hairs slaty; in the 
anal region this colour passes into a ferruginous patch, . . . Skull 
with a very long muzzle, [nearly] equalling that of . . . pyrrhom- 
erus. . . ." 

Type description. — The backs of the ears weakly buffy-white. 
Dorsal surface a dark brindle or grizzle of buffy and black or dark 
gray. Underparts generally dull yellowish, somewhat as in lokriah, 
but much paler. Under side of chin and throat grayish white. Yel- 
lowish of under parts diminished posteriorly and at rear of abdomen 
remains as a mere wash. Insides of fore limbs gray white, hind limbs 


yellowish. Base of tail beneath and anal area dull rusty brown; 
distal part of tail beneath gray with trace of buff. 

Dremomys rufigenis (Blanford) 

Definition. — The subspecies constituting the species rufigenis as 
it is recognized here, are rufigenis, opimus and the extraterritorial 
belfieldi. (See also the synonymy.) The distribution of this species 
is mapped in Figure 26. 

Diagnosis. — The distinguishing characteristics of species rufigenis 
are (1) The pelage of the cheeks is red. (2) The ventral pelage of the 
tail is rich red. (3) There is no rich red color patch in the dorsal 
pelage of the thigh, and the throat is not bright red. 

The above characters distinguish species rufigenis from other spe- 
cies of Dremomys as follows: lokriah by 1 and 2; pernyi by 1 and 2; 
pyrrhomerus by 3; everetti by 1 and 2. 

Intraspecific variation. — Like Dremomys pernyi the present spe- 
cies has in the past been divided rather freely into numerous, faintly 
differentiated subspecies. It is a semi-terrestrial foothills species, 
generally found at less than 5000 feet elevation. Its range centers 
about northern Thailand, but extends south from there into the 
Malay States, north through upper Burma a short distance into 
Assam on the west and eastward into the edge of Yunnan, China. 
It occurs throughout hilly Laos, Tonkin, and Annam, and north a 
little from Tonkin into the adjacent edge of Yunnan. See Figure 26. 

The distinguishing characters of the species rufigenis are two: 
the brilHant colored red pelage of the cheeks, and of a ventral poste- 
rior region including both the anal area and the full-length of the 
undersurface of the tail. The cheeks vary from as pale as Orange 
Rufous to as intense as Dragon's-blood Red, and the other patch 
from Dragon's-blood Red to Brazil Red and occasionally even ap- 
proaching Morocco Red. These marks contrast with the drab, olive- 
gray agouti pelage which rather uniformly cloaks the back and sides 
and upper surfaces of the appendages throughout this genus. Post- 
auricular patches are small and usually white or buffy. A suggestion 
of a black middorsal line has been noted in some material from the 
northwest and southeast extremes of the species range. The ventral 
body pelage is gray at the base and white at the tips. The dorsal tail 
hairs are white-tipped and have a long subapical black band (ca. 10- 
12 mm.) below which is a quite white band less than half as long, and 
a basal portion of less intense black. The dorsal pelage of the feet. 

INDIA / vi 


7\NDj\Ai^ ^E7{^ 




Fig. 26. The species range of the red-cheeked long-nosed squirrel, Dremomys 
rufigenis, as shown by plotting collecting localities of material examined. Sub- 
species: A, rufigenis, B, [extraterritorial] belfieldi; C, opimus. The dotted line 
between the localities for rufigenis and opimus separates these two subspecies as 
known from the localities plotted here, but is not offered as a boundary line between 
the subspecies. 



limbs and hindquarters and also the ventral pelage of the body, are 
subject to infusion with warm buff color in some geographic areas. 

Seven names have been proposed as subspecific variants within 
Dremomys rufigenis as we recognize the species here. We are unable 
to see any justification now for retention of more than three, the 
centrally located typical race, and one each in the northernmost and 
southernmost extremities of the arms of its range extending narrowly 
out from there. 

Dremomys rufigenis rufigenis (Blanford) 

Sciurus rufigenis Blanford, 1878, Jour. Asiatic Soc. Bengal, 47, part 2, p. 156, 

pi. 8 (in color). 
Funambulus rufigenis fuscus Bonhote, 1907, Abstr. Proc. Zool. Soc. London, 

p. 2; 1907, Proc. Zool. Soc. London, 1, p. 10. 
Dremomys rufigenis adamsoni Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 

23, p. 25. 
Dremomys rufigenis ornatus Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, 

p. 26. 
Dremomys rufigenis laomache Thomas, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, 

p. 182. 

Types. — Sciurus rufigenis (lectotype, Thomas, 1921, p. 183) BM 
No., an adult male from above 5000 feet on Mt. Mooleyit, 
Tenasserim, Burma, taken February 17, 1877, by W. T. Blanford; 
fuscus, BM No., an old individual from Bali, 250 meters, 
near Nhatrang, Annam, taken November 10, 1905, by Dr. Vassal; 
adamsoni, BM No., a young male from southeast of Maymyo 
at 2800 feet in Burma, taken July 13, 1913, by J. P. Cook; ornatus, 
BM No., an old female from Yunnan, China, probably 
near Mongtze, collected February 4, 1910, by Alan Owston's col- 
lector H. Orii; laomache, BM No., an old female from "Ban 
Hoi Mak camp," 39 miles southwest of Ti Lao, near Pak Hin Bun, 
Mekong River, Laos, taken on February 29, 1920, by Herbert C. 
Robinson and C. Boden Kloss. 

Material examined, from Assam, Upper Burma, and western Yun- 
nan. — "Animole," Manipur (BM), one; Gokteik, 2133 feet, northern 
Shan States (CNHM), one, (BM), one; Kalaw, 4500 to 4800 feet. 
Southern Shan States (BM), two; Kindat, 250 feet, Chindwin River 
(BM), four; Pyaunggaung, 2794 feet, Northern Shan States, Burma 
(BM), three; Ratnamti, 2000 to 2500 feet. Upper Burma (BM), two; 
Thamaung [Thandaungl, 20 miles E. of Tonjoo [Toungoo], Burma 
(CNHM), one; "Ta-ho," Ind'p't Kareni [Karenni], Burma (USNM), 


Material examined, from Thailand. — Chiengmai [Muang Chiang 
Mai], 700 feet (BM), one; Doi Sritepe [Doi Suthep], 900 meters, 
Chiengmai, Siam (BM), one; Khun Tan [Doi Khun Tan], N. Siam 
(USNM), four; Khun Tan Mts. [Doi Khun Tan], 3000 feet, Siam 
(USNM), one, (CNHM), one; Me Taw Forest, 1200 feet, Raheng, 
Siam (BM), one; Southwest Siam, 14° 20' N., 99° 00' E. (BM), one; 
Doi Hua Mot, Siam (USNM), one; Doi Angina [Doi Ang Ka=Doi 
Inthanon], Siam (USNM), one, (MCZ), 14; Doi Chiengdao [Doi 
Chiang Dao=Doi Luang Chiang Dao=Doi Laem], Siam (USNM), 
two; Doi Sutep [Doi Suthep], Chiengmai, Siam (USNM), one, 
(AMNH), two; Hue Yah Pla [Huai Nua Pla], Siam (USNM), one; 
Doi Lak Sen [Doi Lak Saen], Siam (USNM), one; Doi Pu Kha [Doi 
Phu Kha], Siam (USNM), one; Doi Nangka [Doi Lanka=Khao Pha 
Cho], Siam (USNM), one, (CNHM), one; "Sawan Mtn., Nua, Ban 
Seio," Loei (USNM), one; "Ban Muang Khai," 1000 feet, Ta Li, 
Loei (USNM), five; "Lom Lo Mtn., Ban Maeo, Goksatawn," Dan 
Sai (USNM), 11; "Nam Lang Mtn.," Ban Khok, Naphung, Dan Sai 
(USNM), one; "Ban Na Muang, Na Haeo," Dan Sai (USNM), two. 

Material examined, from Indochina. — Backan [Bac Kan], 500 feet. 
Tonkin (BM), three; Boa-ha, Tonkin (BM), one; Chapa, 5000 feet. 
Tonkin (BM), seven, (MCZ), two, (USNM), two, (CNHM), six; 
Pakha, 1400 meters, Annam [Pa Kha, Tonkin], (CNHM), one; Col 
de Taloun, Laos (MCZ), two; Lo-gui-ho, 5000 feet. Tonkin (BM), 
one; Moung Mo, Tonkin (USNM), one, (CNHM), two; Ngoi Tio, 
Tonkin (BM), one; Phu Kobo, Laos (MCZ), one; Tam-dao, 3000 
feet. Tonkin (BM), three; Lieng San [Leng Sang], Tonkin (CNHM), 
one; Col des Nuages, 400 meters, Annam (BM), five; Moung Moun, 
1200 feet, S. of Lai Chau, Tonkin (CNHM), one; Dak-to, Annam 
(BM), one; Muong Yo, 2300 feet, Laos (CNHM), two; Djiring, 3500 
feet, Annam (BM), one; Kontoun [Kontum], Annam (BM), two; 
Phong Saly, 4400 feet, Laos (CNHM), one; Nape, 2500 to 3000 feet, 
Laos (BM), two; Ban Ton Phung, Laos (USNM), one; Xien Quang- 
Koo, Laos (BM), five; Bali, 250 meters, near Nhatrang, Annam 
(BM), two; Langbian Peak, Annam (BM), one, (MCZ), one, 
(CNHM), 11; Fimnon, S. Annam (USNM), one; 1000 feet above 
Thateng [Ban Thateng, Laos], F.L (CNHM), one; Thua-Luv, 150 
feet, Annam (BM), one. 

Material examined, from Lower Burma. — Ban Sompan, Lower 
Burma (AMNH), one. 

Original description. — "Upper parts dark olive, frizzled, cheeks 
ferruginous, a small white spot behind ear, lower parts white; tail 


hoary, black with white rings in tips above, chestnut below. The 
hairs of lower parts are dark grey at the base, white at the ends, 
there is a tinge of rufous on fore neck and throat in some specimens. 
. . . Tail clad above with black hairs having a white ring near, but 
not at their base, and white tips . . . lower surface of the tail chest- 
nut . . . 

"Skull differs ... in the narrow and singularly elongate nasal por- 
tion. . . . The nearest approach [to rufigenis] is perhaps made by 
S. pernyi . . . [which] has a yellow spot behind the ear. . . . Sciurus 
lokriah also possesses ... a small whitish tuft behind the ear. ..." 
Thus from the very beginning Blanford correctly associated rufigenis 
with pernyi and lokriah. 

The Indian Museum cotype has the dorsal color a grizzle of 
brownish buff and black, the bases of the hairs dark gray. The head 
is more reddish, but the cheeks are clear chestnut without red. Backs 
of ears white, their edges gray. No basiauricular patch. Tail basally 
like the back, terminally darker, composed of hairs with buffy white 
bases, long blackish subterminal rings and short whitish tips. Hands 
and feet and thighs reddish brown. Under parts buffy white with 
gray bases from behind chin to base of tail. The chestnut of the 
cheeks on either side meets beneath the chin. The under side of 
tail from vent as far as observable (tip broken off) is bright rusty red, 
with overall width of 18 mm. Beyond, on either side the black of the 
subterminal rings described for the dorsal surface replaces the red. 

Dremomys rufigenis belfieldi (Bonhote) [Extraterritorial] 

Funambulus rufigenis belfieldi Bonhote, 1908, Jour. Fed. Malay States Mus., 3, 
p. 9, pi. 1 (in color). 

Type.— BM No., adult female from Gunong Ulu Kali, 
Selangor, 4800 to 5800 feet, collected February 9, 1906, by H. C. 

Material examined. — Bukit Fraser, Pahang, Malaya (USNM), 
one; Semangko Pass, Selangor-Pahang boundary, 2500 to 4500 feet 
(BM), two; Gunong Ulu Kali, Selangor, 4800 to 5800 feet (BM), 
two; Kao Tung Sawng [Khao Na Khae], Peninsular Siam, 2500 feet 
(BM), one. 

Original description. — "Differs from the typical rufigenis in hav- 
ing the back paler and slightly grizzled. The hairs are dark at their 
bases with reddish tips, whereas in the typical F. rufigenis they are 
of a much brighter red, which is continued to their base the small 


patch behind the ear . . . bufRsh [in rufigenis], is in this form pure 
snow white. 

"The much redder cheeks and snow-white patch behind the ear 
form very characteristic marks of this race by which it may be easily 

Habits. — Robinson and Kloss (1915, p. 122) report that, "In Se- 
langor this squirrel is confined to the ridges of the higher mountains 
where it lives a partially terrestrial existence among the giant Pan- 
danus and the zerophytic plants clothing the summits. InBandon 
on the other hand it descends the hills and is found on the ground 
amongst the ordinary tropical vegetation of a submontaine forest." 
Their two from Bandon were from above 1200 feet elevation. 

Dremomys rufigenis opimus Thomas and Wroughton 

Dremomys rufigenis opimus Thomas and Wroughton, 1916, Jour. Bombay 
Nat. Hist. Soc, 24, p. 237. 

Type. — BM No., adult male from Hkamti, east bank 
of upper Chindwin River, 500 feet elevation, collected July 25, 1914, 
by G. C. Shortridge and S. A. Macmillan. 

Material examined.— Kaunghein, E. Bank [Chindwin R.], North- 
ern Burma (AMNH), one; Tasu Bum, Northern Burma (AMNH), 
one; Mokokchung, Naga Hills, 5000 feet (BM), two; Nanyaseik, 
Northern Burma (AMNH), one; Hkamti, Upper Chindwin, 500 feet 
(BM), one; Chenga Hka, Northern Burma (AMNH), one; Lahkaw 
Hka, Northern Burma (AMNH), one; Nam Tamai Valley, 4000 feet, 
27° 42' N., 97*' 58' E. (BM), one; Chu-Tun, 8000 feet, Yunnan (BM), 

Original description. — "Like adamsoni . . . but darker and richer 
in colour throughout . . . colour of back more suffused with rufous. 
.... Posterior back and hips suffused with ferruginous instead of 
the lighter and more buffy suffusion of adamsoni. Muzzle deep fer- 
ruginous . . . crown mixed with ferruginous and black, as compared 
with the grey crown of adamsoni. Postauricular patch white, much 
larger and more conspicuous than in [adamsoni]." 

Dremomys pyrrhomerus (Thomas) 

Definition. — The poorly known species pyrrhomerus is composed 
of three very distinct subspecies, pyrrhomerus, riudonensis, and gu- 
laris, and their distribution as known to us from material examined 
is shown in Figure 27. 



n'N+ J. 












/ miNANJ 





1 ..,.,,_( 



Fig. 27. The geographic range of the red-hipped long-nosed squirrel, Dre- 
momys pyrrhomerus, as indicated by plotting collecting localities of material exam- 
ined. Subspecies: A, pyrrhomerus; B, gularis; C, riudonensis. 

Diagnosis. — The diverse species pyrrhomerus possesses the follow- 
ing characteristics of the pelage : (1) There is a red patch in the dorsal 
pelage of the thigh, or a red throat. (2) The ventral pelage of the 
tail is rich red. The first of these two characteristics distinguishes 
pyrrhomerus from rufigenis, and both characteristics distinguish pyr- 
rhomerus from the other three species of Dremomys. 

Relationships to other species. — Zahn (1942, pp. 135-138) recog- 
nizes no species pyrrhomerus but treats pyrrhomerus, riudonensis, 
gularis, and melli as subspecies of D. rufigenis. Allen (1940, pp. 654- 
657) treats the three of these four known to occur in China the same 
way. Our view of pyrrhomerus as a separate species is in agreement 
with that of Osgood (1932, p. 284). The range of D. p. pyrrhomerus 
is separated by a 400-mile hiatus from the nearest parts of the known 
range of rufigenis. This area is not represented well by collections 
of any squirrel species, and this hiatus may or may not be real. We 
have seen 73 specimens of subspecies pyrrhomerus, and its pelage dif- 


ferences from rufigenis are possession of a prominent red thigh patch 
and lack of the red cheek patch. The measurable difference in length 
of snout is considerable, as shown in Figure 28. The difference in 
the appearance of the skulls is greater than that between rufigenis 
and lokriah or between rufigenis and pernyi. The degree of differ- 
ence is, therefore, as exemplied by species within the genus, appar- 
ently one of species rank. 

There are two poorly-known forms located geographically be- 
tween the general range of rufigenis and that of subspecies pyrrhom- 
eru^. One is riudonensis from Hainan and the nearby mainland. We 
have seen only the type series of five specimens of this, from Hainan, 
but we have synonymized the known material of the other poorly- 
known form, melli, from the nearby mainland with riudonensis on the 
basis of the type description. This squirrel has the strong red thigh 
patch of pyrrhomerus, red cheeks like rufigenis but red all over the 
sides and top of the head as well. The small series of broken skulls 
appear to be structurally more like pyrrhomerus than rufigenis. The 
color characters appear superficially to be intermediate between rufi- 
genis and pyrrhomerus but the total relationships appear to be closer 
to pyrrhomerus even though riudonensis is geographically much nearer 
to the known range of rufigenis. We are skeptical, therefore, of the 
existence of a breeding connection between rufigenis and riudonensis. 

□ a a Q a D. lokriah (70) 

ll HIIIin i B B rfFFR Q a o m D. pernyi (65) 


Q • p. gulorit 
3 □ D. pyrrhomerus (42) 

n p H Ft rfti m r 

fTBl Q I I I I l.l.l M l-l I I I I I I 

m*an ' 47.4 

□ a B an mm D. rufigenit (40) 

m*on ' 49.1 

a a B) □ □ cBxD a ffl &n D. •vtrttll (2Q) 

ni«on •44.7 

Fig. 28. Comparison of museum material representing the five species of 
Dremomys in orbitonasal length, presented as percent of total skull length. Arith- 
metic mean for each species is given below the histogram, and the size of the 
sample for each species is indicated in parentheses after the name. The distribu- 
tion of this character in the subspecies D. p. gularis is indicated within that of its 
species by dots. Each square represents one specimen. 



n 29 30 31 32 

□ B R m 1 1 n m NTF mffli B^ a D. lokriah (70) 

mean = 30.1 

S = calidior 
B = howelli 

mean = 27.8 


□ o a i a 

mean = 27.5 



03 a D 

mean = 28.4 

a a [ 

ArffR n 

D m 


D. pernyi (65) 

□ - gularis 
D. pyrrhomerus (42) 

D. rufigenis (40) 

D. everetti (28) 

mean = 28.9 

Fig. 29. Comparison of museum material representing the five species of 
Dremomys in length of orbit expressed as percent of greatest length of skull. 
Arithmetic mean is provided for each histogram, and the size of the sample for 
each species is indicated in parentheses after the name. Each square represents 
a specimen. Where subspecies cluster prominently in the upper or lower range 
for their species in a character, this is shown by special symbols. 

More complicated even than the relationships of riudonensis, are 
those of gularis. We have examined 38 specimens of this form, how- 
ever. This squirrel is known only from the vicinity of Cha Pa, Ton- 
kin, and possibly only from the top of the one mountain, Fan Si Pan. 
Dremomys rufigenis was taken on all sides of this locality, possibly 
always at lower elevations, and there are evidently no intermediate 
specimens. Red cheek and red thigh marks both seem to be present 
but obsolescent in gularis. It possesses the brilliantly red under side 
of the tail in common with both species pyrrhomerus and rufigenis 
but differs from either of them by having the throat and under side 
of the hind legs colored almost as red. These latter characters are 
faintly suggested in occasional individuals of both rufigenis and pyr- 
rhomerus, but are intense and constant characters in the sample of 
gularis. One color character which it shares with pyrrhomerus is a 
faint, blackish middorsal stripe noticeable in about half of the speci- 
mens. This occurs also in one geographic race each of lokriah and 
pernyi but not at all in rufigenis. Osgood (1932) in describing gularis 
as new, noted that the resemblance between skulls of gularis and 


pyrrhomerus is close. We observe that in Figure 29 in proportional 
length of orbit, gularis agrees better with rufigenis, and that in Fig- 
ure 28 in proportional length of snout, gularis lies between but 
perhaps a little nearer pyrrhomerus. In rather intangible general 
appearance and in larger size, gularis is more like pyrrhomerus. Be- 
cause of the altitudinal relationship between gularis and rufigenis, 
and absence of intergrades, they are more likely to be specifically 
distinct than subspecifically. We consider that gularis is best placed 
as a subspecies in the species pyrrhomerus until someone can bring 
new evidence to bear. 

Dremomys pyrrhomerus pyrrhomerus (Thomas) 

Sciums pyrrhomerus Thomas, 1895, Ann. Mag. Nat. Hist. (ser. 6), 16, p. 242. 

Type. — BM No. 2.6.10,66, adult female from Ichang, north bank 
Yangtze River, longitude 111° 20' E., China, collected November, 
1893, by F. W. Styan. 

Material examined, from Szechwan. — 30 miles south of Chunking 
(BM), one; "Huo Chiao Pa" (USNM), two, (CNHM), four; "Yen- 
Ching-Kao," Wanhsien (AMNH), 21, (CNHM), eight; "Tan Kao" 
(CNHM), one; "Yen Tien Pa" (CNHM), two; Yung Cha Shan 
(CNHM), two; Lu Chang Pu (CNHM), six; Pien Ngai (CNHM), 
four, (USNM), one; Wanhsien (MCZ), five. 

Material examined, from Kweichow. — Sunjang [Suiyang] (BM), 
four; Shuan Lung Chang (CNHM), one; Tung Wong Tien (CNHM), 
one; Hsing Liao Pa (CNHM), one; Wenpshui (CNHM), one; Tsunyi 
Hsien (CNHM), one. 

Material examined, from Hupeh. — Chang Yang Hsien (MCZ), 
one; Ichang (BM), four; Tsuk kon Shih [Tung Shih] (MCZ), one. 

Material examined, from Kansu. — "Ma Chu" (AMNH), one. 

Original description. — ". . . Allied to ... S. rufigenis . . . and 
S. pernyi, . . . with both of which it shares the olivaceous back, grey 
and white belly, yellowish postauricular spots, and characteristically 
coloured tail, white-grizzled black above and brilliant red below. 
Sides of cheeks with vague orange suffusion; anal region greyish 
white, like the rest of the under side. A large and prominent blotch 
on the outside of each thigh brilliant rufous. 

"Skull with an enormously elongated muzzle ..." 
Habits.— Walter Granger collected the series from Yen-Ching- 
Kao, Wanhsien during the winters 1921-26. In unpublished field 
notes at the end of 1922-23 he remarks, "The squirrels (5 specimens) 


were all shot at one place, a rough rocky slope with coarse grass and 
some bushes bordering an area where corn is raised in the summer. 
These squirrels are almost entirely terrestrial apparently and live in 
holes in the rocks. During the winter months they come out only oc- 
casionally. . . . The species seems to be of very spotty distribution." 

Dremomys pyrrhomerus riudonensis (J. A. Allen) 

Funambulus riudonensis J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 22, 

p. 472. 
Dremomys melli Matschie, 1922, Beitr. Fauna Sinica, 88, p. 23. 

Types — Funambulus riudonensis, AMNH No. 26651, adult female 
from Riudon, lowlands of Hainan, collected March 11, 1903 by 
A. Owston; Dremomys melli, ZMHU No. 43354, adult male from Yiu 
Shan, Kwangtung, 1000 meters, 230 km. north of Canton, collected 
by R. Mell. 

Material examined. — Riudon, Hainan (AMNH), 4. 

Type description. — The type of riudonensis appears to be in moult, 
as the wool hairs are exposed in the area behind the shoulders. The 
dorsal gray is tinged with reddish in unmoulted areas of the back. 
The head is bright reddish brown, shading to orange red on sides of 
head and cheeks. The hind limbs each have a strong reddish thigh 
mark. Feet and hands are grayish brown, the hands grayer than the 
feet. The ear bases have well-marked white spots. The underparts 
are very pale, whitish gray (the bases of the hairs are gray). The 
underside of the tail is bright rusty red from base to tip. 

Dremomys pyrrhomerus gularis Osgood 

Dremomys pyrrhomerus gularis Osgood, 1932, Field Mus. Nat. Hist. Publ., 
Zool. Ser., 18, p. 284. 

Type.— BM No., an old male from Mt. Fan Si Pan, 
probably above 5000 feet, collected December 3, 1929, by Jean Dela- 
cour and W. Lowe. 

Material examined.— Cha Pa, Tonkin (CNHM), 16, (MCZ), 
three, (USNM), one, (BM), 14; Lo-gui-ho, 5000 to 7000 feet. Ton- 
kin (BM), six. 

Original description. — "Similar to D. pyrrhomerus and D. rufi- 
genis but chin and throat and inner sides of hind legs rich Ochra- 
ceous Tawny in abrupt contrast to other under parts [which are 
bluish, about Clear Payne's Gray]; flank patch obsolescent and re- 
duced to a narrow line scarcely more evident than in rufigenis; 


cheeks, nose and forehead less tawny than in rufigenis, nearly or 
quite as in pyrrhomerus." 

Discussion. — Osgood's further remarks seem well worth quoting 
in part: "This is a very distinct form, . . . the fact that it occurs at 
high elevations within the area occupied by rufigenis lead[s] to the in- 
ference that it is most probably a southern representative of pyrrhom- 
erus. ... In cranial characters, also, the resemblance between the 
two is obviously close. ..." We find that in orbitonasal length the 
sample of eight gularis fall in the lower half of the extremes for this 
measurement known for the species pyrrhomerus, and within the 
range of species rufigenis (see Figure 5) . In orbit length the gularis 
sample might be said to fit better with the rufigenis sample than with 
that of the species pyrrhomerus. However, when compared in series, 
the gularis certainly look more like pyrrhomerus than like rufigenis. 

Dremomys everetti (Thomas) [Extraterritorial] 

Sciurus everetti Thomas, 1890, Ann. Mag. Nat. Hist. (ser. 6), 6, p. 171. 

Type.— BM No., young adult male from Mount Pen- 
risen, west Sarawak, collected in January of 1890 by A. H. Everett. 

Material examined, all from Borneo. — Mt. Kina Balu (BM), 
three, (CNHM), two, (MCZ), 35, (USNM), 19; Mt. Penrisen, Sara- 
wak (NR), one, (BM), two; Sarawak (NR), one; Mt. Tibang 
(MCZ), one. 

Original description. — "Fur thick and soft, . . . Colour uniform 
dark grizzled olive, . . . sides of cheeks, shoulders, and front of hips 
with a very faint fulvous suffusion. Under surface dirty greyish 
white, the hairs everywhere slaty grey for two thirds their length, 
then tipped on the throat and belly with dirty white and on the chin 
and breast with dull fulvous. Ears short, rounded, not tufted or 
emphasized in colour. Tail unusually short, comparatively short- 
haired, almost cylindrical, the hairs ringed with dull fulvous and 
black. Skull small and lightly built, muzzle proportionally very long 
and narrow. . . . Molars small and delicate, their series on the two 
sides parallel. ..." 

Definition. — The species Dremomys everetti is monotypic and en- 
demic to Borneo where it is a montane ground squirrel of the forest. 

Diagnosis. — The species everetti, although conservative, is the 
most distinctive as well as the most isolated species of Dremomys: 
(1) It habitually develops a sagittal crest in the adult. (2) It has 
no whitish tips to the tail hairs. (3) It has no red pelage. (4) It 
has no rich yellow pelage. (5) It has a slight flash mark on the thigh. 


The above characteristics distinguish species everetti from con- 
generic species as follows: from lokriah by 1, 2, 4, and 5; pernyi by 
1, 2, and 5; rufigenis by 1, 2, and 3; and pyrrhomerus by 2 and 3. 

Systematic history. — This species has endured some shifting about 
from one genus to another on the tides of developing opinion and 
knowledge of sciurid relationships in the Oriental Region. It was 
still called Sciurus (Funamhulus) everetti as recently as 1900 by Old- 
field Thomas, Funamhulus everetti as late as 1933 by Banks, and 
Rhinosciurus everetti in 1940 by Allen and Coolidge. Placing of the 
species everetti in the genus Dremomys, however, dates back to 
the 1918 review of the Sciuridae of the Oriental Region by Robinson 
and Kloss. 

Pelage color. — Color notes on stuffed skins of 22 adults of Dre- 
momys everetti at the Museum of Comparative Zoology are offered 
here. Tips of ventral pelage generally are about Cream Color where 
the tips lie dense enough to eclipse the gray proximal color of the 
hairs. This density is most frequent on the throat or in the mid- 
sagittal ventral line. In a few individuals these tips are as pale as 
Light Buff, but in some of the males it is as rich as Light Ochraceous- 
Buff. On the chins the hairs are short and have no dark basal por- 
tion and are consistently Light Ochraceous-Buff. The basal portion 
of the ventral pelage varies among individuals from about Deep 
Neutral Gray to Dark Neutral Gray. The scrotum in eight adult 
males is covered with pelage which is brighter than Ochraceous-Buff 
if not attaining the intensity of Ochraceous-Orange. In one other 
adult male this scrotal pelage is but Warm Buff. The dorsal pelage 
is agouti with general color of Raw Umber in most specimens, but 
one (MCZ No. 36477) is infused with red and is Chestnut-Brown, 
and one or two others are intermediate. The agouti hairs of the 
dorsum seem to have a single light band which is subterminal, and 
are black- tipped. The tail hairs have three or more light bands (as 
many as five), each generally about three millimeters long. One is 
subterminal, and proximal to it is a black band which in most in- 
dividuals is five or more millimeters long. The other dark bands 
separating light ones are shorter and less black. There is a very 
small, generally rather inconspicuous, postauricular patch of Light 
Gull Gray pelage. Legs, feet, and ears are like the back. The tail 
seems faintly annulated from dorsal view in some specimens, partic- 
ularly ones with new tail pelage of less than full length. A slight hip 
mark brightens the dorsal pelage to about Tawny Olive at its junc- 
ture with the ventral pelage on the thigh. The immatures, even as 


young as MCZ No. 36226, which must have been still a nestling feed- 
ing only on milk, are not notably different in color from adults. 

The skull. — Since the four other, more widespread species and not 
this one chanced to be examined by Moore (1959, p. 203) for number 
of transbullar septa and other generic skull characters, it is worth 
mentioning that in the Museum of Comparative Zoology series of 
27 skulls which possess undamaged auditory bullae, all agree with 
the other species in having a single septum in each bulla. This spe- 
cies, everetti, also possesses the other skull characters attributed to 
the subtribe Callosciurina (Moore, 1959, p. 173). Although the tem- 
poral foramen is often small and sometimes absent from one side of 
the skull in other Oriental long-nosed squirrels, it seems to be en- 
tirely absent from skulls of Dremomys everetti. The generic charac- 
ter of Sundasciurus is produced, according to D. Dwight Davis (in 
litt.), by inflation of the musculotubular canal where it leaves the 
auditory bulla anteromesially, and thus constitutes the apparent 
third division of the auditory chamber, between the forks of the 
otherwise single transbullar septum. (This is illustrated by Moore, 
1959, Figure 1, a.) It is interesting that six of the 27 skulls of D. 
everetti have this canal inflated quite like those in some forms of sub- 
genus Aletesciurus. 

Immatures. — To ascertain which individuals should be considered 
adult and therefore more likely to be suitable for taxonomic use, 
comparisons were made within the Museum of Comparative Zoology 
series. Any individual with deciduous fourth upper premolars pres- 
ent or the sagittal suture still partly open between the parietals was 
considered immature. In some of the younger of these the third 
upper molars had not yet erupted to the level of the occlusal plane, 
and in three there were still sutures evident about the interpari- 
etal bone. 

In seven of the eight immatures there was a deciduous third up- 
per premolar present as well as the fourth. Deciduous third upper 
premolars have rarely been commented upon in squirrels. Hall 
(1926, p. 390) has observed this deciduous tooth in the California 
ground squirrel, Spermophilus beecheyi, and comments "the milk 
tooth is only two-thirds the diameter of the permanent tooth." In 
Dremomys everetti the diameter of the deciduous one measures only 
0.2 mm. (with dial calipers under binocular dissecting microscope), 
whereas the permanent third upper premolars for this series range 
from 0.8 to 1.0 mm. The deciduous third upper premolar is a slightly 
curved, slender rod in shape and has a cap of enamel 0.2 mm. or less 


in length. None of these deciduous third premolars reaches the oc- 
clusal plane and only one reaches more than about halfway from 
maxillary to occlusal plane. 

Baculum. — There are a number of bacula preserved with the ma- 
terial of the Museum of Comparative Zoology, and we find that these 
compare much better with the figures which Pocock (1923, p. 223) 
gives of Dremomys than the one he gives of Rhinosciurus. (He shows 
one for a Dremomys dawsoni, incidentally, which may possibly be a 
misreading of bad handwriting for Dremomys adamsoni.) 

Habits. — This is primarily a ground squirrel which lives in the 
deep forest of high mountains in northern and western Borneo, and 
is common between 3000 and 6000 feet elevation but said to be rare 
at 11,000 feet on Mt. Kinabalu. Places from which it has been reli- 
ably reported other than listed above under "material" are Mt. Dulit, 
Mt. Poi, Mt. Trus Madi, the Kelabit Uplands, and Pamambo Range. 
Harrison (1954, p. 162) examined stomachs of 26 from Mt. Trus 
Madi. Five were empty, but from the 21 he found 19 contained in- 
sect material, averaging 35 per cent of the contents per stomach. 
The average amount of fruits and nuts in these was 4 per cent and 
leaves and shoots 15 per cent. The remainder was "other vegetable 
material" which he considered probably included bait. 


Genus MENETES Thomas 

Menetes Thomas, 1908, Jour. Bombay Nat. Hist. Soc, 18, p. 244. 

Type species. — Sciurus berdmorei Blyth. 

Definition. — The genus Menetes is constituted by a single species, 
berdmorei, endemic to the southern part of the Indochinese Subregion 
and a small part of the Malaysian Subregion just south of the Isthmus 
of Kra. See Figure 31. 

Diagnosis. — The genus Menetes has several distinguishing char- 
acteristics: (1) The upper molars and fourth upper premolar possess 
extraordinarily high relief, the paracone with the paraloph and the 
metacone with the metaloph (Bryant, 1945, p. 278) rising high and 
the central valley sinking especially deep between them so that when 
the cusps are planed off by wear, the central valley persists filled with 
dirt and circled by a thin line of enamel. (2) There is one bony sep- 
tum crossing the chamber of the auditory bulla. (3) The baculum 
consists of two separate parts, a shaft and a blade. (4) The length 
of nasal exceeds the least interorbital breadth. (5) The coronoid 
process of the mandible is poorly developed and only faintly falcate. 

Fig. 30. Skull and left mandible of the Indochinese ground squirrel, Menetes 
berdmorei, AMNH No, 54793, Xl. Two left upper molars are missing. 



The above characters distinguish Menetes from other genera of 
the Sciurinae in the Indian and Indochinese subregions as follows: 
from Ratufa by 1, 2, 3, 4, and 5; Funambulus by 1 and 3; Callosciurus 
by 1, 4, and 5; Tamiops by 1, 4, and 5; Dremomys by 1 and 5; and 
Sciurotamias by 1, 2, and 5. 

Systematic history. — Thomas (1914, p. 23) reviewed the species 
berdmorei and recognized five subspecies then constituting the spe- 
cies. In 1918 Robinson and Kloss placed all of the forms then known 
to belong to the genus Menetes in the single species berdmorei. Al- 
though Zahn (1942, p. 104) relegated Menetes to subgeneric rank 
under Lariscus, this was not followed by Ellerman and Morrison- 
Scott (1951), with whom we agree that Menetes is a monotypic genus. 
There remains primarily the question which of the nine races that 
Robinson and Kloss recognized in 1918, the ten listed in 1940 by 
Ellerman, or the six admitted by Ellerman and Morrison-Scott in 
1951 can be sustained. The degree of color variation is very limited. 
The entire genus ranges over a comparatively small area from Mt. 
Popa in central Burma eastward to Annam and Cochin China, and 
southward for a short distance down the Malay Peninsula beyond 
the Isthmus of Kra. In a geographical sense Menetes stops where the 
more southern Lariscus begins. 

Menetes berdmorei (Blyth) 

Definition. — The species berdmorei includes subspecies berdmorei, 
■peninsularis, moerescens, decoratus, consularis and mouhotei, and the 
named forms included in some of these as synonyms. The range of 
the species as known to us from specimens is shown in Figure 31. 

Diagnosis. — Because the genus is monotypic, the diagnosis for the 
genus fits the species as well. 

Intraspecific variation.- — The most apparent variation in the ma- 
terial of this species is in its black dorsal and lateral lines. While 
two whitish longitudinal stripes mark each side of this squirrel with 
great constancy, there may be two or three black stripes on each side 
and one black middorsal stripe, or some or all of the black stripes 
may be missing. 

By plotting the localities and the months from which the extreme 
specimens of the whole species were taken, we find the following: 

(1) The squirrels without any black lines come preponderantly 
from the relatively dry, rain-shadow area of middle and northern 



+ 10"^ 

Ci^lS\^ JEt^ 

Fig. 31. Geographical distribution of the Indochinese ground squirrel, Menetes 
berdmorei, as indicated by plotting localities of material examined. Dotted lines 
separate subspecies on the basis of specimens examined from the plotted localities 
but are only speculative for any other localities. Between consularis and mouhotei 
a considerable area of intergradation is indicated. Subspecies: A, berdmorei; B, pen- 
insularis; C, moerescens; D, decoratus; E, consularis; and F, mouhotei. 

(2) The squirrels with all black lines present come preponderantly 
from mountainous areas which receive the monsoon rains to a greater 

(3) In addition to being distributed according to dry and wet 
geographic areas, these same pattern extremes are evidently varying 


Table 14. Seasonal variation in occurrence of black stripes within 
wet and dry geographic areas. 103 specimens of Menetes berdmorei. 

No black Blackish Black Blackish Black 

lines lateral lateral dorsal dorsal 
Dry Geographic Area 

Rainy Season 7 9 3 3 7 

Dry Season 25 6 1 2 

Wet Geographic Area 
Rainy Season 7 13 

Dry Season 4 3 2 10 1 

to some extent with the wet and dry seasons. In the dry geographic 
area where most individuals without black lines occur, the available 
specimens were taken predominantly in the dry season (25 to 7). 
The majority of those dry area squirrels recorded with some black 
lines were taken in the rainy season in addition to being taken near 
the margin of the dry area. In the wet geographic areas where de- 
velopment of black lines predominates, the few specimens having no 
black lines at all were taken only in the dry season, and those with 
the quite black middorsal lines were taken almost entirely in the 
rainy season (13 to 1) . See Table 14. 

The type of Sciurus berdmorei Blyth is lost. The original descrip- 
tion by Blyth (1849, p. 603) shows the type to be entirely lacking a 
black middorsal line. The describer implied that according to his 
information the type came from the "Thoungyeen district," where 
the available material from Myawadi and nearby Kaukaryit [Kaw- 
kareikl also show lack of a black middorsal line. Was Thomas (1914, 
p. 24) not in error to describe the dry area form as new (M. h. con- 
sularis) and identify the material of Rangoon, Martaban, and the 
neighboring wet coastal area with the type of berdmorei? Blyth's 
description of the type fits the third category in Table 14, i.e., black 
side stripes but no black middorsal. Thomas' interpretation (op. cit., 
p. 23) would put it in the fifth category, i.e., possessing black side 
lines and black middorsal line. Blanford's (1878, p. 162) delineation 
of the Myawadi and Kawkareik material places them in the first and 
second categories of Table 14, i.e., having no black stripes. Only the 
type of berdmorei was, therefore, intermediate between the charac- 
teristic pelage of the subspecies of the coastal area and that of the 
inland subspecies of northern Thailand. It was also more closely 
associated geographically with material which Blyth's description 
identifies with the inland form. Were the designation of the type 


locality less inclusive and vague, this would necessitate applying the 
name M. b. berdmorei exclusively to the subspecies now known as 
M. b. consularis. If the dark-lined coastal form does take its mid- 
dorsal stripe into some part of the Thaungin River Valley, it can be 
argued that Berdmore may (or must) have obtained the type from 
that part. 

See Table 15 for an indication of the body and skull dimensions 
of the species berdmorei. 

Menetes berdmorei berdmorei (Blyth) 

Sciurus berdmorei Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 603. 
Lariscus berdmorei amotus Miller, 1914, Smithsonian Misc. Coll., 61, no. 21, 
p. 24. 

Types. — Sciurus berdmorei (lost), from Thoungyeen District, 
Tenasserim, collected by Captain Berdmore; amotus, USNM No. 
124152, adult male, from Domel Island, Mergui, collected Janu- 
ary 30, 1904, by W. L. Abbott. 

Material examined, from peninsular Thailand. — Klong Ban Lai 
[Ban Salui], Patiyu (BM), two; Koh Lak [Prachuap Khiri Khan], 
sea level (BM), one. 

Material examined, from Burma. — Kau Karyil, Houng Thrau 
[Haung Tharaw] River (BM), one; Bankachon, V.P., south Tenas- 
serim (BM), one; Sullivan's I. (BM), one; Thaget, Little Tenasserim 
River (BM), one; Banlaw, Great Tenasserim River (BM), one; Kis- 
sieraing I. [Kissaraing or Kittha-reng], 50 feet (BM), one. 

Original description. — "Nearly one half larger than [Funambulus] 
palmarum: the prevalent colour grizzled black and golden fulvous, 
with an obscure pale central dorsal streak, flanked by a blackish 
band: this again by a conspicuous yellowish- white line from the 
shoulder to the croup; then blackish again, with a second lateral 
whitish band; below again dusky; and the underparts yellowish 
white, passing to ferruginous towards the vent and underneath the 
tail. Head tinged with ferruginous: . . . This species, according to 
information received from D. F. Lonsdale, inhabits the Thoungyeen 
district [Thoungyin River]," 

Habits. — One collector makes the following field observations: 
"This species spends most of its time on the ground, occasionally 
it may be seen running along railings or up and down slanting or 
broken bamboos, but never at any distance from the ground. At 
Bankachon it is said to be often found on the edges of rice fields, 

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around Maliwun it was occasionally seen running across tracks and 
among long grass, and bamboo scrub, especially in the early eve- 
nings, but I have also seen it in the thickest forest. It is very like 
Tupaia in its movements, hiding at the smallest noise and not read- 
ily making a second appearance. Weight — 73^ ozs." (G. C. Short- 
ridge in Wroughton, 1915b, p. 713) 

Menetes berdmorei peninsularis Kloss 

Menetes berdmorei peninsularis Kloss, 1919, Jour, Nat. Hist. Soc. Siam, 3, 
p. 375. 

Type.— BM No., adult male, from Ban Koh Klap [Ban 
Na], Nakon Sritamarat [Nakhon Si Thammarat], near Bandon, pen- 
insular Thailand, collected July 3, 1913, by H. C. Robinson and 
Cecil B. Kloss. 

Material examined. — Kampenpet [Ban Kamphaeng Phet, latitude 
r 11' N.], Thailand (BM), one. 

Although the type was described as having the median and lat- 
eral black lines strongly marked, it has been previously pointed out 
(Robinson and Kloss, 1915, p. 121) that the Menetes from elsewhere 
(Bandon) are extremely variable in this respect. 

Menetes berdmorei moerescens Thomas 

Menetes berdmorei moerescens Thomas, 1914, Jour. Bombay Nat, Hist. Soc, 23, 

p. 24. 

Type. — BM No. 6.11,6.32, young female from Bali, near Nha- 
trang, latitude 12° N., south Annam, 

Material examined. — Kontoum [also, Kon-toum and Kontum], 
Annam (MNHN), two, (BM), 4; Dak-To [also Dak- to], Annam 
(MNHN), one, (BM), two; Ban Me Thuot [Buon Ma Thuot], An- 
nam (CNHM), four; "Eaktur," E. of Ban Me Thuot, Annam 
(CNHM), two; Ninh Hoa, Annam (CNHM), one. 

Original description. — "Markings about as in decoratus, but less 
conspicuously contrasted, owing to general body colour being darker 
and duller, more olive brown, underside and tips of tail hairs yellow- 
ish. Size rather larger than in other forms. Muzzle of skull unusu- 
ally slender." 

Menetes berdmorei decoratus Thomas 

* Menetes berdmorei decoratus Thomas, 1914, Jour. Bombay Nat, Hist. Soc, 23, 
p. 23. 


Type. — BM No., old female, from Mt. Popa, south cen- 
tral Burma, below 4000 feet, collected April 20, 1913, by G. C. 

Material examined, all from Burma. — Mt. Popa, 4961 feet (BM), 
eight; Mt. Popa (AMNH), five, (CNHM), two (topotypes); Pegu 
Yoma Range, north of Rangoon (AMNH), eight; North Zamayi 
Res., 70 miles north of Pegu, 700 feet (BM), one; South Zamayi Res., 
60 miles north of Pegu, 500 feet (BM), one; 30 miles N.W. of Toun- 
goo, Burma (CNHM), two. 

Original description. — "Median dorsal and upper lateral dark 
streaks prominent, all the markings very strongly defined, the main 
dark lateral band broad and glossy black; an additional blackish 
streak edging the belly; general body colour clear grizzled olive, 
undersurface and tips of tail hairs pure white." 

Discussion. — A glance at the map shows that the Pegu Yoma 
range is a southward extension from the prominent Mt. Popa. Speci- 
mens from these mountains tend to have the black striping heavily 
accentuated. It appears likely that the present race decoratus is 
bounded on the west by the Irrawaddy River and on the east by the 
Salween River. Eastward beyond the Salween the relatively less 
striped race berdmorei occurs. 

Intensification of the black striping recurs elsewhere only in pen- 
insularis and moerescens, both of which are from mountainous regions 
of heavy rainfall. 

Habits. — This form has been observed in the field and commented 
upon as "Occurring on Mt. Popa among rocks and stones, that are 
surrounded by thick scrub, and often close to cultivation, up to 4,000 
feet. Very shy, running into holes and crevices at the slightest sound 
or movement. It is essentially a ground squirrel, seldom, if ever, 
ascending trees, though by no means confining itself to open coun- 
try." (G. C. Shortridge, in Wroughton, 1915a, p. 474) 

Menetes berdmorei consularis Thomas 

Menetes berdmorei consularis Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, 
p. 24. 

Type. — BM No., young female, from Nan [Muang Nan], 
200 meters, Siam, collected October 6, 1901, by T. H. Lyle. 

Material examined, from Thailand. — "Kowpla," Paknampho [Ban 
Pak Nam Pho], Nakonsawan (USNM), one; Airawan Mtn. [Khao 
Erawanl, Lop Buri (USNM), one; Mt. Angka [Doi Inthanon], 4300 
feet [1310 meters] (MCZ), six, (CNHM), one; "central Siam" 


(MCZ), one; Doi Nangkeo [Doi Phi Phan Nam] (MCZ), two; Chien- 
grai [Muang Chiang Rai] 410 meters (BM), three; "Tahkamen" 
[west of Bangkok] (BM), one; halfway between Pichet [Ban Nai 
Muang] and Paknampo [Ban Pak Nam Pho], 33 meters (BM), one; 
100-125 miles north of Bangkok, 20-25 meters (BM), two; "Nam 
Phi" Nan, 225 meters (BM), one; Nakon Sawan [Nakhon Sawan], 
29 meters (BM), one; Me Wong (river), 53 miles east of Um Pang 
[Ban Le Kathe] (BM), three, (AMNH), three; Koon Tan [Doi Kuhn 
Tan, or Sathani Kuhn Tan] (NR), one; Chum Poo [Sathani Tha 
Chomphy] (NR), one; "Vieng Nun," northern Siam (NR), one; 
Muang Prom [Muang Phrom Buri] (BM), two; Me Ping River 
(AMNH), one; below Paknampo [Ban Pak Nam Pho], 26 meters 
(BM), one; Melamoung [Me Lamung] (AMNH), one; north of 
Raheng [Ban Rahaeng] (NM), one; Raheng [Ban Rahaeng], 120 
meters (BM), one; Sokotai [Ban Thank], 64 meters (BM), one; 20 
miles west of Kempanpet [Changwat Kamphaeng Phet] (AMNH), 

Material examined, from Burma. — Taok, Thuaungyin Valley 
[Thaungyin Valley], 1100 feet (BM), one; Myawadi [Mya Wadi], 
Tenasserim (BM), two; Kawkereik [Kawkareik], Tenasserim 
(AMNH), one. 

Original description. — "No median dorsal stripes or upper dark 
lateral ones, the only dark streak being that between the light lat- 
eral streaks, and even this is absent in January and February. Under 
surface yellowish white." 

Menetes berdmorei mouhotei (Gray) 

Sduriis mouhotei Gray, 1861, Proc. Zool. Soc. London, 1861, p. 137. 
Sciurus pyrrhocephalus Milne-Edwards, 1867, Rev. Mag. Zool. (ser. 2), 19, 

p. 225. 
Menetes berdmorei koratensis Gyldenstolpe, 1917, Handl. K. Svenska Vetensk. 

Akad., 57, no. 2, p. 39. 

Types. — Sciurus mouhotei, BM No., from Cambodia; 
pyrrhocephalus, MNHN No. 1864-682, adult female, collected in 
Saigon, Cochin China, by Rudolphe Germain; koratensis, NR No. 4, 
adult male, from Sakerat [Ban Chakkrarat], east Siam, collected 
January 9, 1912, by Nils Gyldenstolpe. 

Material examined, from Thailand. — "Khow Wing," Siracha [Ban 
Si Racha], Chonburi (USNM), one; Satahip (Ban Sattahip], sea level 
(BM), one; Chantabun [Chanthaburi] (USNM), four, (AMNH), four; 


Klong Yai (BM), two, (USNM), three; Lem Ngop [Ban Laem Ngop] 
(BM), three, (USNM), one; Ok Yam [Ok Pyam] (BM), one, (USNM) 
three; Koh [Ko] Kut I. (USNM), six, (BM), six; Koh [Ko] Chang I. 
(BM), one, (USNM), two; Sriracha [Ban Si Racha] (USNM), three, 
(CNHM), one; Hup Bum [Ban Hup Bum], 500 feet (152 meters) 
(BM), two; Ban Sadet [Ban Phan Sadet] near Sriracha (USNM), 
one; Nonokhor [Ban Nong Kho], near Sriracha (USNM), one; "Nong 
Mong," Muong Karbin [Ban Kabin Buri] (USNM), one, (BM), one; 
Kao Sabab [Khao Sa Bap] (USNM), one; Pak Jong [Ban Pak Chong] 
(USNM), one, (AMNH), three, (MCZ), one; Aranya [Ban Aranya- 
prathet] (USNM), one; Lat Bua Kao [Ban Lat Bua Khao] (USNM), 
six; Sakarat [Ban Chakkrarat] (NR), one; 50 miles south of Bangkok, 
seacoast (BM), two. 

Material examined, from Indochina.^ — Plateau Bolovens, Laos 
(AMNH), one; An-Bierh [An Binh] (BM), one; Honquan [Hon 
quan] (BM), two, (CNHM), one; Phu-Quoc I. (BM), one; Tay 
ninh, 100 feet (30 meters) (BM), two; Lagna River (AMNH), one; 
Dinquan [Din Quan] (USNM), one; "Cambodia" (BM), two; Sien- 
Reap [Siem reap] (MNHN), one; Angkor, 150 feet (46 meters) (BM), 
one; Cam Chay [Kam Chay] Mts. (BM), two. 

Original description. — "Grisled grey-brown, with pale rings; lips, 
chin, throat, and under side of body and inside of limbs white; upper 
part of the sides with a longitudinal black streak, edged above and 
below with a narrow white line; tail blackish, whitish washed, hairs 
elongate, brown, with two broad black rings and a white tip. . . ." 

Habits. — Little seems to be known about the natural history of 
Menetes. Gyldenstolpe (1914, p. 15) makes an interesting comment, 
which in his experience appears to apply to this form and consularis. 
"This species was very common in the dry forests both in Northern 
and Eastern Siam. It was always observed near the villages, or in 
the compounds in the towns, but never far into the jungles." 


Sciurotamias Miller, 1901, Proc. Biol. Soc. Washington, 14, p. 23. 
Rupestes Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 398. 

Type species. — Sciurotamias, Sciurus davidianus Milne-Edwards; 
Rupestes, Rupestes forresti Thomas. 

Definition. — The genus Sciurotamias is constituted by the two 
monotypic subgenera Sciurotamias and Rupestes, both endemic to 
northern China. 

Diagnosis. — (1) There are two dorsal processes on the zygomatic 
process of the squamosal. (2) The ectopterygoid ridge of the ali- 
sphenoid is obsolescent. (3) The upper incisors are strongly opistho- 
dont, (4) Three bony septa cross the chamber of the auditory bulla. 

The above characters all distinguish Sciurotamias from each of 
the other genera of the Sciurinae of the Indian and Indochinese sub- 

Subgenus SCIUROTAMIAS Miller 

Sciurotamias Miller, 1901, Proc. Biol. Soc. Wachington, 14, p. 23 

Type species. — Sciurus davidianus Milne-Edwards. 

Definition. — The subgenus Sciurotamias includes only the type 
species davidianus, a polytypic, montane, ground squirrel of north- 
ern China. 

Diagnosis. — The subgenus Sciurotamias apparently may differ 
from subgenus Rupestes in several notable characters: (1) The squa- 
mosal is high. (2) The zygoma ascends to a point well above halfway 
on the height of the rostrum. (3) The superior process of the jugal 
is very low and anterior to the midlength of the orbit. (4) There is 
no pronounced temporal ridge. (5) The postorbital process of the 
frontal is more than 2 mm. long. (6) There is a peg-shaped third 
upper premolar. (7) There are generally faint whitish postauricular 
streaks on the pelage but no whitish longitudinal stripes on the sides 
of the body. 




Fig. 32. Skull and left mandible of the montane rock squirrel of China, Sci- 
urotamias davidianus, AMNH No. 45370, X 1. Note indications of the three trans- 
bullar septa which distinguish this genus from the Oriental tribe Callosciurini and 
the Holarctic tribe Sciurini as well. 

Sciurotamias davidianus Milne-Edwards 

Definition. — The species davidianus is composed of subspecies 
davidianus and consobrinus, wiiich include several named forms here 
as synonyms. The geographic distribution known to us from speci- 
mens examined is shown in Figure 33. 

Diagnosis. — Since the subgenus is monotypic, the diagnosis of 
the species is identical to that for the subgenus. 

Relationships to other species. — Sciurotamias is said to inhabit cliffs 
and rocky ground of the mountains avoiding trees, and nesting in 
deep crevices. It would be interesting to know the extent to which 
it differs from Dremomys p. pyrrhomerus in habitat. The two seem 
not to have been collected at the same localities; although their 
ranges appear to be closely approximated. G. M. Allen (1940, p. 646) 
has stated that Dremomys is arboreal in contrast to Sciurotamias, 
but he evidently inferred this from Dremomys being taken in deep 
forest. References to its habits in Malaya and Borneo indicate that 




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Fig. 33. Geographic distribution of the Chinese rock squirrel, Sciurotamias 
davidianus (round symbols), and Chinese cliff squirrel, Sciurotamias (Rupestes) 
forresti (square symbols), as revealed by plotting the collecting localities of mate- 
rial examined. Subspecies of species davidianus: A, davidianus; B, consobrinus. 

it inhabits forested areas of mountains, but is primarily terrestrial 
there. In 1934, at a single locality in the mountains 25 miles west 
of Wenchwan, Szechwan, T. Donald Carter collected Dremomys 
pernyi and Sciurotamias d. consobrinus. Since this is the only place 
which we have located where both of these species are known to 
have been collected, his comments on their habitats seem especially 
significant. He recalls clearly (September 1956) in conversation over 
the collected material laid before him, that the Dremomys pernyi in- 
habited the steep, forested slope of the narrow valley of Chengou 
Creek, and he recalled them particularly in the second growth near 


the foot of the slope where there had been some removal of timber. 
They did ascend trees, he said, but seemed to be on the ground a 
good deal, and he collected them on the ground, particularly among 
moss-covered logs of down timber. The Sciurotamias lived in small 
areas of rather level, boulder-strewn grassland where the narrow 
valley bottom widened out occasionally. There were no trees on 
this, and Sciurotamias lived here like a ground squirrel. The tree- 
less character of these level areas along the creek bottom Mr. Carter 
attributed to perhaps seasonal flooding by the torrential mountain 

The evidence presented in the accompanying maps shows that 
the ranges of both Dremomys pernyi and Dremomys pyrrhomerus lie 
to the south of the distribution of Sciurotamias davidianus (excepting 
for the isolated example of the latter from Hwangtsaopa, Kweichow) . 
The above example of apparently sharp segregation into two very 
different biotic communities in an area of overlap of species range, 
may not, however, be a fair indication of the variety of habitat uti- 
lized by either species in the parts of its range which are not shared 
with closely related species. 

It may be worth noting in this connection that the distribution 
of Sciurotamias davidianus shown here coincides with the long north- 
east to southwest portion (but not the southeast) of the "Deciduous 
broad-leaved forest predominated by deciduous oaks" mapped by 
Wang (1956, p. 513) and that that of S.forresti lies in an area in which 
Wang shows two alternating vegetation types: 1. Tundra and alpine 
vegetation, and 2. Montane coniferous forest predominated by spruce 
and fir. Dremomys p. pyrrhomerus occurs south of this in the area 
that Wang maps as "Evergreen broad-leaved forest of evergreen 
oaks, shima, and laurels, with [pines] in secondary stands." D. p. 
pernyi is sympatric with S. forresti but apparently allopatric with 
S. davidianus. D. p. flavior is allopatric with D. p. pyrrhomerus but 
occupies a more western part of the evergreen broad-leaved forest 
area as mapped by Wang. 

Although Sciurotamias is as long-snouted proportionately as Dre- 
momys lokriah, for example, a skull of Sciurotamias is readily distin- 
guished from those of Dremomys species by being broader, arched, 
and flattened, so that it is shaped quite like skulls of the chipmunk, 
Eutamias. The posterior upper premolar is notably smaller than 
any of the upper molars in Sciurotamias whereas in Dremomys, its 
size does approximate that of one or more of the molars. The fora- 
men ovale is circular in Sciurotamias, narrowly oval in Dremomys. 


Sciurotamias has three bony septa across the chamber of the auditory 
bulla; whereas members of the tribe Callosciurini have one or none. 
The upper incisors are strongly opisthodont in Sciurotamias; where- 
as those of members of the Callosciurini are orthodont or, more fre- 
quently, proodont (defined in Moore, 1959, p. 162). Sciurotamias 
differs also in having cheek pouches. 

See Table 15 for an indication of the body and skull dimensions 
of species davidianvs. 

Sciurotamias davidianus davidianus (Milne-Edwards) 

Sciurus davidianus Milne-Edwards, 1867, Rev. et Mag. Zool. (ser. 2), 19, 

p. 196. 
Dremomys latro Heude, 1898, Mem. Hist. Nat. Chinois, 4, pt. 2, pp. 54-55. 

Types. — Sciurus davidianus, MNHN No. 1863-655, collected by 
Pere Armand David in the "mountains of Pekin," Hopei, China; 
latro, not seen, but thought to be from Shantung, China (Allen, 1940, 
p. 662). 

Material examined, from Hopeh, China (except as noted). — "Chi- 
hli" (NR), 15; near Ching Wang Tas [Chinwangtao] (USNM), 11; 
Hsin-lung-shan, 65 miles N.E. of Peking [Jehol, Manchurial (USNM), 
two; Wu-ting-shan, 75 miles N.E. of Pekin [Jehol, Manchuria] 
(USNM), two; Western Hills, 15 miles W. of Peking (USNM), one, 
(RNH), one; Tung-Hng (USNM), one, (AMNH), three, (CNHM), 
four; Eastern Tombs (AMNH), three, (CNHM), one. 

Material examined, from Shansi, China. — Mts. 50 miles N.W. of 
Tai-yuan-fu [Yangku] (USNM), four; Mts. 10 miles S. of Wu-tsai 
[Wutai] (USNM), one; Mts. 70 miles N.N.W. of Tai-yuan-fu [Yang- 
ku] (USNM), three; "He-shuin" (MCZ), one; "Sjol" (NR), five. 

This subspecies is distinguished from the southern race by its 
lighter, grayer dorsal color, about Deep Grayish Olive, and by its 
grayish instead of buffy-brownish under parts. The race is appare- 
ently confined to territory north and east of the Hwang-Ho River. 
The postauricular patches appear to be less prominent in this race, 
and the posterior streaking from them, mentioned above, is evident 
less frequently. 

We follow Allen (1940, p. 662) in synonymizing latro with da- 

Sciurotamias davidianus consobrinus (Milne-Edwards) 

Sciurus consobrinus Milne-Edwards, 1868-74, Recherches pour servir a I'Hist. 
Nat. des Mammiferes, p. 305. 


Dremomys collaris Heude, 1898, Mem. concern. I'Hist. Nat. de I'Emp. Chinois, 

4, part 2, p. 55, pi. 12, figs. 2-2c. 
Dremomys saltitans Heude, 1898, Mem. concern. I'Hist. Nat. de I'Emp. Chinois, 

4, part 2, p. 55, pi. 12, figs. 4-4c, 
Sciurotamias owstoni J. A. Allen, 1909, Bull. Amer. Mus. Nat. Hist., 26, p. 428. 
Sciurotamias davidianus thayeri G. M. Allen, 1913, Mem. Mus. Comp. Zool., 

40, no. 4, p. 231. 

Types. — Sciurus consobrinus, not found at MNHN, a specimen 
from Muping, Szechwan, China, collected by Pere Armand David; 
collaris, not seen, possibly at the Sikawei Museum in Shanghai, 
China; saltitans, not seen, possibly at the Sikawei Museum in Shang- 
hai, China, thought to have been taken in Hupeh Province, China 
(Allen, 1940, p. 665) ; owstoni, AMNH No. 27545, an adult female 
from Si-Tai-pa-shieng Mts., Shensi, China, taken in October 1905 by 
A. Owston; thayeri, MCZ No. 8008, an adult male from Washan, 6000 
feet, Szechwan, China, taken May 17, 1908, by Walter R. Zappey. 

Material examined, from Shensi, China.^ — Tai-Pa-Shiang Mts., 
6000 feet (AMNH), eight; Tai-pei-shan Dist., 3000 feet, 80 miles 
W.S.W. of Sian-fu [Siking] (USNM), three; base of Tai Pei Shan, 
Tsingling Mts. (CNHM), six; 15 miles east of Yen-an-fu Fushih 
(USNM), one; Tsingling Mts. (AMNH), three, (MCZ), one, 
(CNHM), six; 45 miles south of Fensiangfu, 3000 to 3600 feet 
(AMNH), two, (MCZ), one, (CNHM), two; nine and 15 miles south 
of Ching-chien-hsien, 2600 feet (USNM), two; Liu-tsuen, 1500 feet, 
15 miles south of Sian-fu (USNM), four. 

Material examined, from Kansu, China. — Ha Tebbuland, 8500 
feet, Wantsangku (MCZ), three. 

Material examined, from Hupeh, China. — Ma Fu Ling, 5000 feet 
(MCZ), two; Hsien Shan Hsien, 4000 feet (MCZ), two; "Tan Sweo 
Nah," 3000 feet (MCZ), one; Fongshan [Fanghsien] (MCZ), one; 
"Moo Swei Ping" (MCZ), one. 

Material examined, from Honan, China. — "Honan" (NR), three. 

Material examined, from Szechwan, China. — "Lu Din Chiao," 
5000 feet (USNM), one; Longpan [Sungpan] (USNM), one; near 
Weichow [Weikiu], 9000 feet (USNM), one; "Tu-pa-keo," Moupine 
[Muping] (CNHM), one; Wenchuan [Wench wan] (USNM), 19, 
(AMNH), 24; Mt. Omei (USNM), two; E. of Romitchangu (MCZ), 
one; Ko-chia-ho-pa, 7500 feet (CNHM), one; Trashi-cho-ten, 8250 
feet (CNHM), one; WaShan (MCZ), one; "Nai Su Chen" (CNHM), 
one; "Goan Shih Dwe" (CNHM), four; "Sungpan" (CNHM), two. 

Material examined, from Kweichow, China. — Whang Tsao Pa 
[Hwangtsaopa] (USNM), one. 


Discussion. — We have been unable to distinguish any really good 
characters to separate consobrinus, owstoni and thayeri from one an- 
other. Taken together they are darker and more brownish, Cinna- 
mon Brown to Mummy Brown, than the northern race davidianus, 
and their under parts have a wash of Light Ochraceous Buff to 
Ochraceous Buff which is lacking in the gray-bellied davidianus. In 
the series of consobrinus from Wenchwan, the white postauricular 
patches are prominent. 

It is rather remarkable that a specimen should come from very 
southern Kweichow over 300 miles south of other Sciurotamias d. 
consobrinus locations found. In this hiatus, moreover, are a number 
of localities where Dremomys pernyi and D. pyrrhomerus were taken. 

Habits. — Fu (1936, p. 258) makes an interesting comment on the 
habits of this squirrel in western Honan where he collected it in the 
Sung-shan: "This animal nests in the ground, and is ascribed as a 
nuisance to peasants on account of its damage on corns. The den 
and its contents are often destroyed by ploughing." 

Subgenus RUPESTES Thomas 
Rupestes Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 398. 
Type species. — Rupestes forresti Thomas. 

Definition.— The subgenus Rupestes includes only the one species 
forresti, which is a relict known only from the mountains of western 
China as shown in Figure 33. 

Diagnosis. — (This diagnosis is based upon one skull of a subadult 
with the occiput and bullae missing, USNM 255138, and four skins.) 
On the evidence examined Rupestes differs from subgenus Sciuro- 
tamias in several notable characters: (1) The squamosal reaches up 
the side of the cranium less than halfway from the base of the zygo- 
matic process to the base of the postorbital process. (2) The anterior 
edge of the zygoma ascends to a point only halfway in the height of 
the rostrum. (3) The superior process of the jugal is high and pos- 
terior to the midlength of the orbit. (4) There are pronounced tem- 
poral ridges. (5) The postorbital process of the frontal is very short 
(less than 1 mm. in this specimen). (6) There are no third upper 
premolars. (7) There is a rather faint light stripe along the flank 
between shoulder and hip. 

Sciurotamias (Rupestes) forresti (Thomas) 

Rupestes forresti Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 399. 


Type. — BM No., old female, from Mekong-Yangtze 
divide, latitude 27° 20' N., 7000 to 9000 feet, northern Yunnan, 
China, collected June 5, 1921 by George Forrest. 

Material examined. — E. of Yalung [River], 15,500 feet [4725 me- 
ters], Szechwan [Sikang] (USNM), one; Chienchwan [Tienchiian], 
Szechwan [Sikang] (MCZ), one; LiChiang [Likiang], Yunnan 
(AMNH), one, (BM), one; Mekong-Yangtze divide latitude 27° 
20' N., 7000 to 9000 feet, Yunnan (BM), two. 

Definition. — This relict, rarely taken species is apparently mono- 
typic, and its range as known from four specimens examined is shown 
in Figure 33. 

Diagnosis. — Since the subgenus is monotypic, the diagnosis of the 
subgenus serves for the species. 

This species is readily told from davidianus by the presence of a 
pair of faint whitish lines high on the sides which extend from the 
shoulders to the hips. These lines are edged on each side by the dark 
grizzled dorsal coloring. Some 6 mm. beyond them the dark dorsal 
color changes to the light orange brown of the sides and that in turn 
to the buffy of the underparts. 

See Table 15 for some dimensions of the type. 


Twenty-six species of the seven genera of Sciurinae occurring in 
the Indian Subregion and Indochinese Subregion of the Oriental Re- 
gion are, for these geographic areas, here fully revised. Since four 
of the seven genera (Funambulus, Tamiops, Menetes, and Sciuro- 
tamias) are endemic, complete generic revisions of these are pre- 
sented. In order to offer a complete revision of the genus Dremomys, 
also, the one extraterritorial (monotypic, Malaysian) species, everetti, 
is included here. Eight species of Callosciurus are recognized in the 
Indochinese Subregion, and are shown to constitute two superspecies 
which are broadly sympatric throughout much of the subregion. 
These two superspecies are observed to form a phylogenetic unit 
separate from the Malaysian species of Callosciurus, and a complete 
revision is offered for this eight species unit. In the genus Ratufa 
complete revision is presented for the two species in the Indian Sub- 
region. The species Ratufa bicolor which occurs throughout much 
of the Indochinese Subregion is revised only for this area, although 
it also has an extensive distribution in the Malaysian Subregion. 

The principal changes from earlier existing concepts are within 
the Indochinese subregional unit of the genus Callosciurus, a unit 
previously unrevised. Considerable advancement in clarification 
and improvement of previously existing knowledge, however, is pro- 
vided for every species by mapping the distribution of collecting 
localities. Future supplementation and correction of the present 
knowledge is made easy by presenting latitudes and longitudes for 
the collecting localities presumed found. In those subspecies for 
which adequate material was examined, particularly in American 
museums, descriptions are provided from large samples in effort to 
make them applicable to the populations (not merely the type speci- 
mens) involved. In the genus Callosciurus in particular, perhaps 
only because more collecting sites and materials are known, striking 
evidence is present in the taxonomic account of inability to cross the 
main courses of the great rivers of the subregion. The evolutionary 
implications of this and other facts of geographic distribution of 
diurnal squirrels in this subregion are only sparingly discussed. 



It is intended in any discussions in the present paper to employ 
the geographic names current at the time of final preparation of the 
manuscript. In the lists citing localities where specimens were col- 
lected, however, the names and spellings that were recorded on the 
original specimen tag by the collector have ordinarily been retained. 
Should any question arise, this permits reinterpretation of the local- 
ity without resort to the original source. The currency of geographic 
names of the smaller principalities actually used here necessarily 
varies a great deal, depending upon the source. For some localities 
no knowledge is available to us except that they are villages near 
which specimens were taken, and they are mapped in an account of 
a collecting trip. Many, however, were taken by collectors whose 
routes and collecting localities were not published and whose local- 
ties to be plotted must usually be found on some map. Since villages 
in the jungle are often moved, localities of 50 years ago may be in 
many cases more easily found on maps made about that time. 

Many sources were utilized, but those most helpful and employed 
most were: Stanford (1917) "Complete Atlas of China" with maps 
at scale of 1:3 million; a set of maps of Indochina and most of Thai- 
land at 1:1/2 million published at Luang Prabang in 1931 by le 
Service Geographique de I'lndochine; "The Times Survey Atlas and 
Gazetteer of the World," J. G. Bartholomew (1922); its successor, 
"The Times Atlas of the World, Mid-Century Edition," John Bar- 
tholomew (1958) ; and maps published by the National Geographic 
Society, Washington, D. C. 

Because villages are moved and the names of towns, cities, and 
even natural geographic features, have been changed by new political 
regimes, latitude and longitude (from the Greenwich zero meridian, 
NOT the Paris one of the map of Indochina cited above) are em- 
ployed. Even this effort at accuracy embraces errors of five miles or 
so from differences of latitude or longitude between different maps. 
This may be a matter of some concern in the immediate vicinity of a 
large river which is potentially an important, long-time barrier to 
the species. 



In alphabetizing, place names of more than one word are treated 
as if but a single word (e.g., Ban Don, Bangnara, Ban Huai Som 
would come properly in this order) , 

Occasionally place names are used that may be found on a good 
map in several places (e.g., Khao Soi Dao is the name of at least 
three mountains in Thailand) . With the knowledge from the speci- 
men tag of who collected the specimen and when, and with available 
knowledge of the collector's itinerary, we have generally been able 
to decide which must be the correct place. In the gazetteer, only if 
specimens here studied were collected at two localities of the same 
name is an effort made to warn the user of the existence of other like 
place names. 

Adung Valley, Burma: 28° 10' N., 97° 40' E. 

Agia, Assam, India: 26° 06' N., 90° 34' E, 

AiNGGYl, Pakokku Chin Hills, Burma: 21° 05' N., 94° 22' E. 

AlRAWAN, Mt.: see Khao Erawan 

Akhe Triangle, Upper Burma: 26° 54' N., 98° 12' E. 

Akola, Berar, India: 20° 45' N., 77° 00' E. 

Amarakantak, India: 22° 40' N., 81° 45' E. 

Amherst, Tenasserim, Burma: 16° 00' N„ 97° 38' E. 

Amlekhganj, Nepal: 27° 18' N., 85° 00' E. 

Amraoti, Bastar [Berar], India: 20° 55' N., 77° 50' E. 

An Binh, Cochin China, Viet Nam: 10° 30' N., 106° 43' E. 

Angkor, Cambodia: 13° 25' N., 103° 50' E. 

Angtong [Ang Thong]: see Ban Bang Kaeo 

Aranya: see Ban Aranyaprathet 

Arbre Broye, Viet Nam: 11° 57' N., 108° 28' E. 

Avalanche, Nilgiri Hills, India: 11° 10' N., 76° 35' E. 

Ayuthia: see Phra Nakhon Si Ayutthaya 

Bag Kan, Tonkin, Viet Nam: 22° 10' N., 105° 45' E. 

Bag Tan Tray, Tonkin, Viet Nam: 22° 06' N., 103° 15' E. 

Baksa, Formosa, China: 23° 10' N., 120° 29' E. 

Balapalli, Madras, India: 13° 45' N., 79° 35' E. 

Bali: see Nhatrang 

Bamanigaon, Assam, India: 26° 10' N., 91° 25' E. 

Ba Nam Chi, Tonkin, Viet Nam: 22° 07' N., 102° 56' E. 

Ba Nam Nhung, Tonkin, Viet Nam: 22° 07' N., 103° 00' E. 

Ban Aranyaprathet, Prachin Buri, Thailand: 13° 40' N., 102° 30' E. 

Ban Bang Kaeo, Ang Thong, Thailand: 14°35'N.,100°25'E. 

Ban Chakkrarat, Nakhon Ratchasima, Thailand: 15° 00' N., 102° 25' E. 

Ban Chang Thuk, Thailand: 14° 45' N., 101° 30' E. 

Ban Chiang Dao, Thailand: 19° 20' N., 99° 00' E. 

Ban Choeng Doi, Chiang Mai, Thailand: 18° 50' N., 99° 10' E. 

Ban Den, Nong Khai, Thailand: circa 18° 00' N., 102° 55' E. 

Ban Doi Saket : see Ban Choeng Doi 

Bandon: see Ban Makham Tia 


Ban Dong: see Ban Pong 

Bang Hue Hom, Thailand: 17° 53' N., 100° 06' E. 

Bang Hue Pong: see Sathani Pang Hua Phong 

Bangkok (160 miles north of), Thailand: 16° 00' N., 100° 45' E. 

Bangkok: see Krung Thep 

Bangnara: see Narathiwat 

Ban Hai Huai Som: see Ban Huai Som 

Ban Hin Laem, Kanchanaburi, Thailand: 14° 40' N., 98° 40' E. 

Ban Hin Ngom, Nong Khai, Thailand: 17° 55' N., 102° 50' E. 

Ban Hoi Mak: see Pak Hin Bun 

Ban Houei Sai, Laos: 20° 18' N., 100° 25' E. 

Ban Huai Som, Nan, Thailand: 18° 20' N., 100° 30' E. 

Ban Huang Som, Trat, Thailand: 11° 50' N., 102° 50' E. 

Banhuathanon: see Ban Hua Thanon 

Ban Hua Thanon, Kamphaeng Phet, Thailand: circa 16° 14' N., 99° 35' E. 

Ban Hu Kwang, Nakon Sawan, Thailand: 15° 55' N., 100° 00' E. 

Ban Hup Bon, Thailand: 13° 05' N., 101° 05' E. 

Ban Kabin Buri, Prachin Buri, Thailand: 14° 00' N., 101° 45' E. 

Bankachon, Tenasserim, Burma: 10° 9' N., 98° 36' E. 

Ban Kang: see Ban Mae Klang 

Ban Khana, Nan Province, Thailand: 19° 19' N., 100° 56' E. 

Ban Khlong Bang Lai, Patiyu, Thailand: 10° 45' N., 99° 10' E. 

Ban Khlong Khlung, Kamphaeng Phet, Thailand: 16° 10' N., 99° 45' E. 

Ban Khlong Mon, Samut Sakhon, Thailand: 13° 35' N., 100° 35' E. 

Ban Khlua Klang, Prachuap Khiri Khan, Thailand: circa 11° 35' N., 99° 39' E. 

Ban Khok, Naphung, Dansai, Loei, Thailand: 16° 25' N., 101° 25' E. 

Ban Khok Klap: see Ban Na 

Ban Khung Samphao, Thailand: 15° 20' N., 100° 00' E. 

Ban Kiriwong: see Ban Wat Khiriwong 

Ban Klua Klang: see Ban Khlua Klang 

Ban Krat, Thailand: 14° 26' N., 103° 04' E. 

Banlad: see Ban Lat 

Ban Laem Ngop, Trat, Thailand: 12° 08' N., 102° 35' E. 

Ban Lat, Phu Kheio, Chaiyaphum, Thailand: 16° 30' N., 101° 55' E. 

Ban Lat Bua Khao, Nakhon Ratchasima, Thailand: 14° 50' N., 101° 35' E. 

Ban Le Kathe, Tak, Thailand: 15° 50' N., 98° 50' E. 

Ban Luang, Chiang Mai, Thailand: 18° 25' N., 98° 40' E. 

Ban Mae Klang, Thailand: 18° 30' N., 98° 40' E. 

Ban Mae Mo, Thailand: 18° 15' N., 99° 45' E. 

Ban Mae Sariang, Mae Hong Son, Thailand: 18° 10' N., 97° 55' E. 

Ban Mae Tan Nua, Lampeng, Thailand: 18° 20' N., 99° 25' E. 

Ban Mae Ton: see Pang Mae Ton 

Ban Maha Chai, Samut Sakhon, Thailand: 13° 30' N., 100° 14' E. 

Ban Makham Tia, Thailand: 9° 10' N., 99° 20' E. 

Ban Manao Wan, Thailand: 14° 55' N., 101° 05' E. 

Ban Manaswan: see Ban Manao Wan 

Ban Meh Na, Thailand: 19° 12' N., 98° 52' E. 

Ban Mb Thuot: see Buon Ma Thuot 

Ban Mi Chai, Nongkhai, Thailand: 17° 50' N., 102° 50' E. 

Ban Muak Lek, Sara Buri, Thailand: 14° 40' N., 101° 10' E. 


Ban Muang Khai, Tha Li, Muang Loei, Thailand: 17° 35' N., 101° 20' E. 

Ban Mung Ky: see Ban Muang Khai 

Ban Nai Muang, Phichit, Thailand: 16° 25' N., 100° 20' E. 

Ban Na, Thailand: 8° 53' N., 99° 17' E. 

Ban Na Ko, Nan, Thailand: circa 19° 10' N., 100° 54' E. 

Ban Nam Chut Yai, Ranong, Thailand: 10° 25' N., 98° 45' E. 

"Ban Na Muang, Na Haeo," Dan Sai, Loei, Thailand: 17° 15' N., 101° 02' E. 

Ban Nam Yen, Koksathon, Thailand: 17° 10' N., 101° 05' E. 

"Ban Na Nong," Chuempae [Ban Chum Phae], Khon Kaen, Thailand: 16° 34' N., 

102° 03' E. 
Ban Neng Kho: see Ban Nong Kho 

Ban Nong Bua, Pasak River, Thailand: 14° 50' N., 101° 05' E. 
Ban Nong Chik: see Ban Tuyong 

Ban Nong Don Ta, Thailand: circa 18° 12' N., 104° 05' E. 
Ban Nong Kho, Chon Buri, Thailand: 13° 10' N., 101° 05' E. 
Ban Nong Pla Lai, Thailand: 16° 05' N., 98° 45' E. 
Ban Non Tao Lek, near Chaiyaphum, Thailand: 16° 27' N., 101° 50' E. 
Ban Non Toulek: see Ban Non Tao Lek 
Ban Pak Chan, Thailand: 10° 32' N., 98° 51' E. 

Ban Pak Chong, Nakhon Ratchasima, Thailand: 14° 40' N., 101° 25' E. 
Ban Pakli: see Ban Sopli 

Ban Pak Nam Pho, Nakhon Sawan, Thailand: 15°'40' N., 100° 10' E. 
Ban Pak Tho, Rajburi, Thailand: 13° 21' N., 99° 50' E. 
Ban Phan Sadet, Thailand: 13° 05' N., 101° 05' E. 
Ban Phone, Laos: 15° 25' N., 106° 35' E. 
Banphotphisai: see Ban Hu Kwang 

Ban Phu Khieo, Chaiyaphum, Thailand: 16° 34' N., 101° 52' E. 
Ban Pong, Rat Buri, Thailand: 13° 50' N., 99° 55' E. 
Ban Pua, Nan, Thailand: 19° 11' N., 100° 53' E. 
Ban Rahaeng, Tak, Thailand: 16° 50' N., 99° 05' E. 
Ban Sadet: see Ban Phan Sadet 
Ban Sahng Kaw: see Ban Sawan Ko 
Ban Sai Kan: see Nawngchik 

Ban Sai Yok, Kanchanaburi, Thailand: 14° 29' N., 98° 50' E. 
Ban Sa Kaeo, Prachin Buri, Thailand: 13° 50' N., 102° 05' E. 
Ban Salok Bat, Kamphaeng Phet, Thailand: 16° 00' N., 99° 45' E. 
Ban Salui: see Ban Khlong Bang Lai 
Ban San Tha, Nan, Thailand: circa 18° 21' N., 100° 40' E. 
Ban Sattahip, Chon Buri, Thailand: 12° 40' N., 100° 55' E. 
Ban Sawan Ko, Sakon Nakhon, Thailand: 17° 27' N., 104° 05' E. 
Ban Sichon, Thailand: 9° 00' N., 99° 55' E. 

Ban Si Khiu, Nakhon Ratchasima, Thailand: 14° 55' N., 101° 45' E. 
Ban Si Racha, Chon Buri, Thailand: 13° 10' N., 101° 05' E. 
Ban Sob Pa [Soppa], Burma: 18° 48' N., 97° 27' E. 
Ban Sompan, Lower Burma: circa 10° 30' N., 98° 30' E. 
Ban Sopli, Thailand: 18° 00' N., 100° 30' E. 
Ban Talok Bat: see Ban Salok Bat 
Ban Tarn Dam: see Ban Than Dam 
Ban Ta Yai: see Ban Tha Yai 
Ban Tayun, Plateau Bolovens, Laos: 15° 23' N., 106° 23' E. 


Ban Tha Chang, Thailand: 14° 35' N., 101° 25' E. 

Ban Tha Khanum, Kanchanaburi, Thailand: 14° 45' N., 98° 40' E. 

Ban Than Dam: Near Ban Si Racha 

Ban Thani, Sukhothai, Thailand: 17° 00' N., 99° 51' E. 

Ban Tha San, Ranong, Thailand: 10° 30' N., 98° 55' E. 

Ban Thateng, Laos: 15° 25' N., 106° 23' E. 

Ban Tha Yai, Thailand: 8° 23' N., 99° 52' E. 

Bantion: see Ban Tayun 

Ban Ton Phung, Laos: 20° 18' N., 100° 05' E. 

Ban Tuyong, Thailand: 06° 50' N., 100° 00' E. 

Ban Um Phang: see Ban Le Kathe 

Ban Wang Mai Khon, Sukhothai, Thailand: 17° 19' N., 99° 50' E. 

Ban Wang Takhian: see Kanchanaburi 

Ban Wat Khiriwong, Thailand: circa 8° 23' N., 99° 52' E. 

Ban Wiang Tai, Mae Hong Son, Thailand: 19° 20' N., 98° 25' E. 

Ban Wichian, Thailand: 15° 40' N„ 101° 05' E. 

Bag Ha, Tonkin, Viet Nam: 22° 10' N., 104° 23' E. 

Barapani: see Borpani 

Baria, Cochin China, Viet Nam: 10° 30' N., 107° 10' E. 

Baurang, Yunnan [Sikang], China: 28° 52' N., 101° 23' E. 

"Bhadwar," Kangra, Punjab: 32° 05' N., 76° 18' E. 

Bhor Phloy Nong Yungchang: see King Bo Phloi 

Bhuket: see Koh Phuket 

BiEN HOA, Cochin China: 10° 52' N., 106° 50' E. 

Bintenna: see Pintenne 

BOK Pyin, Tenasserim, Burma: 11° 15' N., 98° 46' E. 

Boraphet, Thailand: 15° 43' N., 100° 14' E. 

BORI, Hoshangabad, India: 20° 55' N., 79° 05' E. 

Borpani, Khasi Hills, Assam, India: 25° 39' N., 91° 53' E. 

Borploy: see King Bo Phloi 

BOUN Tai: see Bun Tai 

Boyce's Point, Tenasserim, Burma: 10° 45' N., 98° 29' E. 

BUKET: see Koh Phuket 

Bung Borapet: see Boraphet 

Bun Tai, Laos: 21° 18' N., 102° 00' E. 

BUON Ma Thuot, Viet Nam: 12° 40' N., 108° 05' E. 

Burnihat, Assam, India: 25° 37' N., 91° 50' E. 

Cachar [Silchar], Assam, India: 24° 42' N., 92° 50' E. 

Cachar Hills, Assam, India: 25° 00' N., 93° 00' E. 

Chaduquet Point: see Laem Set Kuat 

Chaiyaphum, Thailand: 16° 27' N., 101° 50' E. 

Chamaphum: see Ban Phu Kheio 

Champang, Tenasserim, Burma: 10° 12' N., 98° 27' E. 

Chamrajnagar, Mysore, India: 11° 55' N., 76° 55' E. 

Chance Island: see Ko Chan 

Chandragiri Pass, Nepal: 27° 39' N., 85° 08' E. 

Changwat Kamphaeng Phet, Kamphaeng Phet, Thailand: 16° 30' N., 99" 30' E. 

Changwat Mae Hong Son, Thailand: 19° 15' N., 97° 55' E. 

Chang Yang Hsien, Hupeh, China: 30° 25' N., 111° 13' E. 

Chantabun: see Chanthaburi 


Chanthaburi, Thailand: 12° 35' N., 102° 05' E. 
Chan Tuk: see Ban Chan Thuk 
Chao Chu, Yunnan, China: 25° 35' N., 100° 18' E. 
Chapa: see Cha Pa 

Cha Pa, Tonkin, Viet Nam: 22° 20' N., 103° 50' E. 
Chaung, 20 miles n.n.w. of Kindat, Burma: 23° 45' N., 94° 17' E. 
Chaung-lung, Salween ferry, Yunnan, China: 24° 12' N., 98° 58' E. 
Chenga Hka, Burma: 26° 07' N., 95° 52' E. 

Chengou Creek, 25 mi. W. of Wenchwan, Szechwan, China: 31° 20' N., 103° 00' E. 
Chenkang, Yunnan, China: 24° 06' N., 99° 22' E. 
Che Pei, Fu Min District: see Fuminhsien 
Cherrapunji, Assam, India: 25° 13' N., 91° 47' E. 
Chettiri Range, Salem Dist., Madras, India: 11° 50' N., 78° 25' E. 
Chiang, Szechwan, China: 28° 47' N., 102° 53' E. 
Chiang Rai: see Muang Chiang Rai 

Chiang Saen Kao, Chiang Rai, Thailand: 20° 15' N., 100° 05' E. 
Chiang Saen Luang: see Chiang Saen Kao 
Cheinat: see Muang Chainat 
Chienchwan: see Tienchiian 
Chieng Dao: see Ban Chiang Dao 
Chiengmai: see Muang Chiang Mai 
Chih ping: see Shihpingchow 

"Chi li ping," Fanghsien, Hupeh, China: 32° 03' N., 110° 54' E. 
Chimeli Pass, Upper Burma: 26° 08' N., 98° 40' E. 
Chinbyit, Burma: 22° 03' N., 94° 40' E. 
Ching-chien-shien: see Tsingkien 

Ching Feng Ling, Fukien, China: 27° 11' N., 118° 13' E. 
Chinteh: see Tsingteh 

Chinwangtao, Hopeh, China: 39° 56' N., 119° 37' E. 
Chin Wang Tas: see Chinwangtao 
Chip-Chip, Formosa: circa 23° 55' N., 120° 45' E. 
Chipwi, northeastern Burma: 25° 52' N., 98° 07' E. 
Chomtong: see Ban Luang 

Chora, Tonkin, Viet Nam: 22° 18' N., 105° 52' E. 

Chulungapu, Upper Tebbuland, Kansu, China: circa 34° N., 101° 30' E. 
Chumphon, Thailand: 10° 30' N., 99° 10' E. 
Chum Poo: see Sathani Tha Chomphu 
Chunganhsien, Fukien, China: 28° 02' N., 117° 53' E. 
Chungan Hsien: see Chunganhsien 
Chung Chiao Miao: see Chiang 

Chung King, Szechwan, China: 29° 34' N., 106° 35' E. 
Chungtang, Sikkim, India: 27° 37' N., 88° 38' E. 
Chuntown: see Chumphon 

Clara Island, Mergui Archipelago, Burma: 10° 50' N., 97° 55' E. 
CoiMBATORE, Madras, India: 10° 55' N., 76° 58' E. 
Col des Nuages, Annam, Viet Nam: 16° 11' N., 108° 10' E. 
Col de Taloun, Laos: 19° 50' N., 102° 15' E. 
COORG, India: 12° 00' N., 76° 00' E. 

"Cotengady Estate," [Nelliampathi Hills, Palghat District], Cochin, India: 
10° 40' N., 76° 35' E. 


CuCHAi: see Kuchai 

Da Ban, Phanrang Province, Annam: 12° 38' N., 109° 06' E, 

Dacca, East Pakistan: 23° 43' N., 90° 26' E. 

Dagung Hka, Burma: 26° 05' N., 95° 55' E. 

Dak-to, Annam, Viet Nam: 14° 40' N., 107° 45' E. 

Daingmhu, 40 mi. N. of Pegu, Burma: 17° 42' N., 96° 14' E. 

Dalat, Annam, Viet Nam: 11° 58' N., 108° 25' E. 

Dalu, N. Burma: 26° 20' N., 96° 12' E. 

Dao Soi Dao: see Khao Soi Dao 

Darjeeling, Bengal, India: 27° 00' N., 88° 18' E. 

Darugiri, Garo Hills, Assam, India: 25° 37' N., 90° 46' E. 

DeLisle, Thailand: 09° 40' N., 98° 20' E. 

Den Chai, near Bang Hue Hom, Thailand: circa 17° 53' N., 100° 06' E. 

Dening, Mishmi Hills, Assam, India: 28° 01' N., 96° 10' E. 

Devikop, S. Mahratta, India: 15° 08' N., 75° 00' E. 

DiKCHU, Sikkim, India: 27° 25' N., 88° 35' E. 

Djiring, Annam, Viet Nam: 11° 35' N., 108° 06' E. 

Doi Angka: see Doi Inthanon 

Doi Chiang Dao: see Doi Luang Chiang Dao 

Doi Chiengdao: see Doi Luang Chiang Dao 

Doi Hua Mot: see Chiengmai 

Doi Inthanon, Chiang Mai, Thailand: 18° 32' N., 98° 32' E. 

Doi Kang Ma: see Doi Khang Ma 

Doi Khang Ma, Chiang Mai, Thailand: circa 18° 08' N., 98° 27' E. 

Doi Khun Tan, Lamphun, Thailand: 18° 30' N., 99° 20' E. 

Doi Lak Saen, Thailand: circa 18° 08' N., 98° 07' E. 

Doi Lak Sen: see Doi Lak Saen 

Doi Langka: see Khao Pha Cho 

Doi Luang Chiang Dao, Chiang Mai, Thailand: 19° 25' N., 98° 55' E. 

Doi Mae Kong Ka, Mae Hong Son, Thailand: circa 18° 06' N., 97° 49' E. 

Doi Me Kong Kha: see Doi Mae Kong Ka 

Doi Mae Lai, Mae Hong Son, Thailand: circa 18° 09' N., 98° 00' E. 

Doi Mei Lai: see Doi Mae Lai 

Doi Nangka: see Khao Pha Cho 

Doi Nang Keo: see Doi Phi Phan Nam 

Doi Par Sakeng, northwestern Thailand: 19° 35' N., 99° 03' E. 

Doi Phi Phan Nam, Chiang Mai, Thailand: 19° 05' N., 99° 25' E. 

Doi Phra Chao: see Khao Pha Cho 

Doi Phu Het, Nan, Thailand: circa 18° 20' N., 100° 35' E. 

Doi Phu Ka, Nan, Thailand: 19° 05' N., 101° 05' E. 

Doi Phu Kha, Thailand: 19° 05' N., 101° 05' E. 

Doi Pu Het: see Doi Phu Het 

Doi Pu Kha: see Doi Phu Kha 

Doi San Huai Wai, Nan, Thailand: circa 18° 20' N., 100° 35' E. 

Doi Sritepe: see Doi Suthep 

Doi Sutep: see Doi Suthep 

Doi Suthep, Chiang Mai, Thailand: 18° 50' N., 98° 55' E. 

DoMEL Island, Mergui, Burma: 11° 37' N„ 98° 15' E. 

Donghai: see Dong Hoi 

Dong Hoi, Annam, Viet Nam: 17° 32' N., 106° 35' E. 


Don Qua, Laos: 19° 45' N., 102° 35' E. 

DOONSA, near junction Gyn [Gyaing] and Haung Thrau [Haung Tharaw] Rivers, 

near Moulmein, Burma; 16° 37' N., 98° 00' E. 
Drachuapchiri Khan Bankhluaklang: see Ban Khlua Klang 
Dreyi, Mishmi Hills, Tibet: 28° 03' N., 96° 15' E. 
DuRAGlRl: see Darugiri 

"Eaktur," E. of Ban Me Thuot: see Buon Ma Thuot 
Eastern Tombs, Hopeh, China: 40° 00' N., 117° 00' E. 
Etawah United Provinces, India: 26° 50' N., 79° 00' E. 
Fanghsien, Hupeh, China: 32° 03' N., 110° 54' E. 
Fansipan: see Mt. Fan-si-pan 

Fengsiangfu, Shensi, China: 34° 24' N., 107° 29' E. 
Feng Yang, Yunnan, China: 24° 45' N., 102° 40' E. 
FiMNON, S. Annam, Viet Nam: 11° 40' N., 108° 22' E. 
Fi Shan Kwan, Szechwan, China: 30° 03' N., 103° 05' E. 
FOOCHOW, Fukien, China: 26° 05' N., 119° 18' E. 
Fuching: see Futsing 

FUCHOW, Fukien, China: 26° 05' N., 119° 18' E. 
Fu Fu Su, Szechwan, China: 29° 35' N., 103° 35' E. 
Fu Lin, Szechwan, China: 29° 20' N., 102° 45' E. 
FUMINHSIEN, Yunnan, China: 25° 12' N., 102° 31' E. 
Futsing, Fukien, China: 25° 40' N., 119° 25' E. 
Gammaduwa, Ceylon: 07° 35' N., 80° 41' E. 
Gangfang, Burma: 26° 06' N., 98° 38' E. 
Gangtok, Sikkim, India: 27° 20' N., 88° 30' E. 
Ghatmatha, Satara District, Bombay, India: 17° 25' N., 73° 40' E. 
GOALPARA, Assam, India: 26° 10' N., 90° 37' E. 
Godavari, Nepal: 27° 35' N., 85° 20' E. 
GoDAVERi: see Godavari 

GoKTEiK, Northern Shan States, Burma: 22° 20' N., 96° 52' E. 
GOLAGHAT, Assam, India: 26° 30' N., 94° 01' E. 
Gopaldhara, Rungbong Valley, India: 26° 55' N., 88° 12' E. 
Gorkha, Nepal: see Satthar 
Gougah: see Lien Gongah 

GuNGWADORiA, north slope of Palni Hills, India: circa 10° 20' N., 77° 30' E. 
Gyaing, and Haung Thrau [Haung Tharaw] Rivers, near Moulmein, Burma: 

16° 37' N., 98° 00' E. 
Ha-Gang: see Ha Giang 

Ha Giang, Tonkin, Viet Nam: 22° 50' N., 104° 58' E. 
Hahti, Burma: 26° 04' N., 95° 32' E. 
Hai Bum, Burma: 26° 02' N., 95° 47' E. 

Hai Yen Hsien, Hangchow Bay, Chekiang: 30° 17' N., 120° 10' E. 
Hambantota, Ceylon: 06° 07' N., 81° 07' E. 

Hastings Island, Mergui Archipelago, Burma: 10° 05' N., 98° 20' E. 
Hathiban, Nepal: 27° 37' N., 85° 15' E. 
Hathiben: see Hathiban 
Hataura, Nepal: 27° 25' N., 85° 05' E. 
Hat Sanuk, Rajburi, Thailand: 11° 52' N., 99° 42' E. 
Haungtharaw River (where crossed by Moulmein-Kawkareik road), Burma: 

16° 30' N., 98° 05' E. 


Heinsum, Burma: 25° 52' N., 95° 32' E. 

Helwak, Satara Dist., Bombay, India: 17° 20' N., 73° 40' E. 

Hetora: see Hataura 

HiNLAEM: see Ban H in Laem 

HiN Lap: see Sathani Hin Lap 

Hkamkawn, Burma: 26° 02' N., 98° 23' E. 

Hkamti, Burma: 26° 00' N., 95° 42' E. 

Hkamti Plain, Burma (taken to be the same locality as Putao and Ft. Hertz): 

27° 22' N., 97° 27' E. 
Hkawni, Gaw, Upper Burma: 25° 58' N., 98° 00' E. 
Hmawbi, 40 km. north of Rangoon, Burma: 17° 05' N., 96° 05' E. 
Hnathial, Lushai Hills, Assam, India: circa 23° 45' N., 92° 55' E. 
Hoi Xuan, Annam, Viet Nam: 20° 20' N., 105° 05' E. 
HOKOW, Szechwan, China: 30° 01' N., 101° 08' E. 
HOMALIN, Burma: 24° 53' N., 94° 55' E. 
HOMU-SHU Pass, Yunnan, China: 24° 55' N., 98° 45' E. 
HOOPBON: see Ban Hup Bon 
HOSHANGABAD, India: 22° 45' N., 77° 45' E. 
HouE SAi: see Ban Houei Sai 

Hpimaw, Salween-Irrawaddy Divide, Burma: 25° 58' N., 98° 38' E. 
HsiAO-MENG-TUNG: see Siaomengtung 
Hsiao Yang Chi, Szechwan, China: 28° 47' N., 102° 44' E. 
Hsien Shan Hsien, Hupeh, China: 31° 30' N., 110° 55' E. 
Hsing Liao Pa, Kweichow, China: 28° 14' N., 106° 33' E. 
HsiN Kai: see Sinkai 

HsiN LUNG SHAN, Jehol, Manchuria: 40° 38' N., 117° 17' E. 
HsiPAW, Shan States, Burma: 22° 38' N., 97° 20' E. 
Htawgaw, Burma: 25° 55' N., 98° 25' E. 
Htingwan, Burma: 26° 31' N., 97° 52' E. 
Huai Nua Pla, Tak, Thailand: 16° 54' N., 98° 48' E. 
Hue Sai, near Koh Lak, Thailand: circa 11° 48' N., 99° 47' E. 
Hue Yah Pla: see Huai Nua Pla 
HuEY Yang: see Ban Si Racha 
HULAUNG, Burma: 25° 10' N., 95° 05' E. 

HUNGAI, 40 miles E. of Talifu, Yunnan, China: circa 25° 30' N., 100° 20' E. 
Hung Fa Chao: see Shanghai 
Hup Bon: see Ban Hup Bon 
Hup Bum: see Ban Hup Bon 
Hwang Tsao Pa: see Hwangtsaopa 
HWANGTSAOPA, Kweichow, China: 24° 57' N., 104° 36' E. 
Ichang, Hupeh: 30° 45' N., 111° 22' E. 
ICHANG Hsien, Hupeh, China: 30° 42' N., 111° 17' E. 
I-Lan Hsien, I-Lan, Taipei, Taiwan, China: 24° 25' N., 121° 42' E. 
Imaw Bum: see Mt. Imaw Bum 
INTHA, Burma: 24° 30' N., 94° 42' E. 

IPIN, Szechwan- Yunnan border, China: 28° 43' N., 104° 32' E. 
Isle [He] de la Table, Tonkin, Viet Nam: 20° 57' N., 107° 30' E. 
James Island, Mergui Archipelago, Burma: 10° 25' N., 98° 17' E. 
Jhimpir Lake, Sind, India: 25° 00' N., 68° 00' E. 
Kabaw Valley, 20 mi. W. of Kindat, Burma: 23° 40' N., 94° 12' E. 


Kachek, Hainan, China: 19° 14' N., 110° 27' E. 

Kadrangyang, Fort Hertz Rd., 75 mi. N. of Myitkyina, Burma: circa 26° 07' N., 

Kagi Dist., Formosa: 23° 29' N., 120° 17' E. 
Kaing R., S. Pyinwana, S. Burma: 19° 44' N., 96° 09' E. 
Kalewa, Burma: 23° 12' N., 94° 13' E. 
Kalhatti [Falls], Nilgiri Hills, India: 11° 27' N., 76° 43' E. 
Kalutara, Matugama, Ceylon: 06° 34' N., 79° 58' E. 
Kampeng Pet: see Kamphaeng Phet 
Kamphaeng Phet, Thailand: 16° 30' N., 99° 30' E. 
Kampot, Cambodia: 10° 40' N., 104° 18' E. 
Kan Bouri : see Kanchanaburi 
Kanchanaburi, Thailand: 14° 00' N., 99° 30' E. 
Kangpu: see Kanpu 
Kangra, Punjab: 32° 05' N., 76° 18' E. 
Kanjanaburi: see Kanchanaburi 

Kanpu, near Hangchow, Chekiang, China: 30° 20' N., 120° 50' E. 
Kao Chiao, Yunnan, China: 25° 01' N., 102° 20' E. 
Kao Hao, Quangtri Province, Annam, Viet Nam: 16° 45' N., 107° 11' E. 
Kao Lem: see Khao Laem 
Kao Luang, Huey Yang: see Khao Luang, Sathani Huai Yang (not the same 

locality as Khao Luang, Nakhon Si Thammarat) 
Kao Sabab: see Khao Sa Bap 
Kao Seming: see Khao Saming 

Kaoshanshih, near Foochow, Fukien, China: 25° 25' N., 119° 28' E. 
Kao Soi Dao: see Khao Soi Dao 
Kaotien, Fukien, China: 27° 18' N., 117° 25' E. 
Kao Tung Sawng: see Khao Thung Song 
Karjine: see Ban Pak Tho 

Karong, Manipur, Assam, India: 25° 15' N., 94° 05' E. 
Katha, Burma: 24° 10' N., 96° 20' E. 
Kauktaung, Burma: 25° 43' N., 95° 25' E. 
Kaunghein, Burma: 25° 42' N., 95° 24' E. 
Kawai, Burma: 26° 02' N., 95° 34' E. 
Kawkareik, Burma: 16° 28' N., 98° 16' E. 
Kawlaw, Southern Shan States, Burma: 20° 38' N., 96° 34' E. 
Kawlichaung, Burma: circa 16° 15' N., 98° 25' E. 
Kaw Nom Plu: see Khao Nam Pliu 
Kellengode: see Kollangod 
Kempanpet: see Changwat Kamphaeng Phet 
Khad Tham Phra, Laos: 20° 15' N., 100° 05' E. 
Khajjean: see Kangra 

Kha Nu, Kampeng Phet, Thailand: 16° 07' N., 99° 46' E. 
Khao Erawan, Lop Buri, Thailand: circa 14° 50' N., 100° 35' E. 
Khao Khat, Nakhon Sawan, Thailand: 15° 43' N., 100° 07' E. 
Khao Kop, Nakhon Sawan, Thailand: 15° 43' N., 100° 08' E. 
Khao Laem, Thailand: 14° 25' N., 101° 30' E. 
Khao Luang, Sathani Huai Yang, Thailand: 11° 36' N., 99° 41' E. (not the same 

locality as Khao Luang, Nakhon Si Thammarat) 
Khao Na Khae: see Khao Thung Song 


Khao Nam Pliu, Thailand: 7° 35' N., 99° 50' E. 

Khao Pha Cho, Chiang Mai, Thailand: 19° 00' N., 99° 25' E. 

Khao Phra Phut, Lop Buri, Thailand: 14° 50' N., 100° 40' E. 

Khao Sa Bap, Chanthaburi, Thailand: 12° 35' N., 102° 15' E. 

Khao Saming, Trat, Thailand: 12° 21' N., 102° 27' E. 

Khao Soi Dao, S.E. Thailand: 15° 30' N., 100° 45' E. (not the same locality as 

Khao Soi Dao, Trong, Thailand) 
Khao Thung Song, Peninsular Thailand: 8° 20' N., 99° 48' E. 
Kharagpur, Bengal, India: 22° 20' N., 87° 20' E. 
Khet Dong Heing, Laos: 17° 53' N., 101° 34' E. 
Khlong Wang Hip, Thailand: circa 8° 12' N., 99° 43' E. 
Khon Kaen, Thailand: 16° 25' N., 102° 40' E. 
Khun Tan Mts. : see Doi Khun Tan 
KiATiNG: see Loshan 

Kienyang, Fukien, China: 27° 21' N., 118° 05' E. 
KiL: see Kotajiri 

Kin, Burma: 22° 41' N., 94° 41' E. 
KiNDAT, Burma: 23° 40' N., 94° 22' E. 

King Bo Phloi, Kanchanaburi, Thailand: 14° 48' N., 99° 10' E. 
King Chiang Saen: see Chiang Saen Kao 

King Island, Mergui Archipelago, Burma: 12° 30' N., 98° 22' E. 
KissERiANG Island, Burma: 11° 41' N., 98° 28' E. 
Klong Ban Lai: see Ban Khlong Bang Lai 
Klong Klung: see Ban Khlong Khlung 
Klong Menao: see Ban Huang Som 
Klong Morn: see Ban Khlong Mon 
Klong Wang Hip: see Khlong Wang Hip 
Klong Yai, Thailand: 11° 44' N., 102° 52' E. 
Ko Chan, Mergui Archipelago, Thailand: 9° 25' N., 97° 50' E. 
Ko Chang, Trat, Thailand: 12° 00' N., 102° 30' E. 
KODAIKANAL, India: 10° 12' N., 77° 30' E. 
KODURA, Balapalli Range, Madras, India: 13° 57' N., 79° 32' E. 
KoH Chang Island: see Ko Chang 

Ko-chia-ho-pa, Szechwan, China: circa 30° 20' N., 102° 20' E. 
KOHIMA, Assam, India: 25° 40' N., 94° 12' E. 
KoH KuT Island: see Ko Kut 
KOH Lak: see Prachuap Khiri Khan 
KoH Lan: see Ko Lan 
KoH Pangan: see Ko Phangan 
KoH Phai: see Ko Phai 

KOH Phuket, Southwestern Thailand: 7° 53' N., 98° 24' E. 
KoH Si Chang: see Ko Si Chang 
KoH Tao: see Ko Tao 
KOKKOAING, Burma: 22° 28' N., 96° 18' E. 
Kokkogon: see Kokkogwa 
KOKKOGWA, Burma: 19° 59' N., 95° 30' E. 
Ko Kut, Trat, Thailand 11° 40' N., 102° 35' E. 
Ko Lak: see Prachuap Khiri Khan 
Ko LAN, Chon Buri, Thailand: 12° 55' N., 100° 45' E. 
KOLLANGOD, India: 10° 38' N., 76° 37' E. 


KOLLEGAL, Coimbatore, India: 12° 10' N., 77° 10' E. 

Ko Maphrao, Thailand: 7° 55' N., 98° 25' E. 

Kontoum: see Kontum 

KONTUM, Annam, Viet Nam: 14° 25' N., 107° 55' E. 

KOON Tan: see Doi Kuhn Tan 

Ko Phai, Chonburi, Thailand: 12° 55' N., 100° 40' E. 

Ko Phangan, Thailand: 09° 45' N., 100° 00' E. 

Ko Si Chang, Chon Buri, Thailand: 13° 10' N., 100° 50' E. 

KOTAJIRI, Nilgiri Hills, India: 11° 25' N., 76° 55' E. 

Ko Tao, Surat Thani, Thailand: 10° 05' N., 99° 50' E. 

KOTTAWA, Ceylon: 06° 03' N., 80° 20' E. 

Kowjeen: see Ban Pak Tho 

KOWKAT: see Khao Khat 

Kowkob: see Khao Kop 

Krabin: see Ban Kabin Buri 

Krat: see Ban Krat 

Kratt: see Ban Krat 

Krong Ba, S. Annam, Viet Nam: 14° 00' N., 108° 18' E. 

Krung Thep, Phra Nakhon, Thailand: 13° 45' N., 100° 30' E. 

KUCHAI, Yunnan, China: 27° 20' N., 103° 14' E. 

Kucheng, Fukien, China: 26° 40' N., 118° 43' E. 

Ku-LU, Szechwan, China: 28° 05' N., 101° 12' E. 

KUMBUKKAN, Uva Prov., Ceylon: 06° 47' N., 81° 17' E. 

Kunming : see Yunnanfu 

KURUMBAPATTI, India: 11° 45' N., 78° 15' E. 

KusHAN: see Kaoshanshih 

Kutung, Yunnan, China: 25° 27' N., 99° 30' E. 

Kwatum: see Kaotien 

KwE Koi: see Mae Nam Khwae Noi 

Kyaukmyaung, Burma: 22° 30' N., 95° 57' E. 

Kyundaw, Burma: 21° 10' N., 94° 40' E. 

Lachen, Sikkim, India: 27° 45' N., 88° 32' E. 

Lachung, Sikkim, India: 27° 43' N., 88° 44' E. 

Lacon Sritamarat: see Nakhon Si Thammarat 

Laem Set Kuat, Thailand: 10° 05' N., 98° 40' E. 

Laem Sing, Chantaburi, Thailand: 12° 30' N., 102° 05' E. 

Lahkaw Hka, Burma: 26° 25' N., 96° 12' E. 

Lahore, Punjab: 31° 35' N., 74° 15' E. 

Lai Chau, Tonkin, Viet Nam: 21° 52' N., 103° 10' E. 

Laitlynkot, Khasi Hills, Assam, India: 25° 27' N., 91° 51' E. 

Lai Yoke: see Ban Sai Yok 

Lakchang Ga, Burma: 26° 16' N., 96° 07' E. 

Lakhon: see Muang Lampang 

Lakya, Tenasserim, Burma: circa 16° 15' N., 98° 25' E. 

Lamaya, Szechwan, China: 29° 52' N., 99° 54' E. 

Lampha, Burma: 16° 18' N., 98° 19' E. 

Lam-ra: see Sathani Lamphura 

Landa [Londa], India: 14° 30' N., 74° 25' E. 

Langbian Peaks, Annam, Viet Nam: 12° to 12° 15' N., 108° 20' E. 

Lang Son, Tonkin, Viet Nam: 21° 50' N., 106° 45' E. 


Langyang, Burma: 25° 54' N., 98° 18' E. 

Lao Bao, Laos: 16° 37' N., 106° 34' E. 

Lao Fou Tchay, Laos: 21° 43' N., 102° 25' E. 

Lao Kay, Tonkin, Viet Nam: 22° 30' N., 104° 00' E. 

Lat Bua Kao: see Ban Lat Bua Khao 

Laukhaung, Burma: 25° 53' N., 98° 13' E. 

Lawksawk, Burma: 21° 13' N., 96° 50' E. 

Lay Song Hong: see Thale Song Hong 

LE BOKOR, Cambodia: 10° 40' N., 104° 00' E. 

Lei-Mui-Mon, Hainan, China: 19° 00' N., 109° 30' E. 

Lem Ngop: see Ban Laem Ngop 

Lem Sing Mountain: see Laem Sing 

Leng Sang, Tonkin, Viet Nam: 22° 25' N., 103° 17' E. 

Lian-feng-Kiang, Omei Shan, Szechwan, China: 29° 25' N., 103° 25' E. 

LiCHlANG : see Likiang 

Lien Gongah, Annam, Viet Nam: 11° 30' N., 108° 20' E. 

Lieng San: see Leng Sang 

Likiang, Yunnan, China: 27° 04' N., 100° 12' E. 

LiMPA, Burma: 25° 45' N., 95° 30' E. 

Lingtam: see Singtam 

Litien, Yunnan, China: 27° 14' N., 99° 27' E. 

Liu-TSUEN, Shensi, China: 34° 02' N., 108° 55' E. 

Lo-Gui-Ho, Tonkin, Viet Nam: 22° 19' N., 103° 42' E. 

LOMLOE Mt., Ban Maeo, Goksatawn, Dahnsai, Loey, Thailand: 17° 00' N., 

101° 00' E. 
Long Lung: see Ban Nong Pla Lai 
LONKIN, Burma: 25° 40' N., 96° 22' E. 
Lonpan: see Sungpan 

LosHAN, Szechwan, China: 29° 35' N., 103° 40' E. 
Lo Tiao, Laos: 20° 20' N., 100° 05' E. 
Lo Tu Kow: see Lukou 
Lsien: see Wutingchow 

Luang Prabang, Laos: 19° 53' N., 102° 10' E. 
Lu Chang Pu, Szechwan, China: 29° 23' N., 108° 10' E. 
LuiA, Chaibassa, India: 22° 35' N., 85° 25' E, 
Lukou, Yunnan, China: 25° 55' N., 98° 50' E. 
LUNGCHOW, Kwangsi, China: 22° 22' N., 106° 54' E. 
Lunghkang (Ga), Burma: 26° 36' N., 97° 48' E. 
LUNGKAi, Yunnan, China: 25° 57' N., 101° 54' E. 
LUNGTAN, China: 32° 05' N., 119° 20' E. 
Lung Van: see Van Lung 
LUTINGKIAO, China: 29° 54' N., 102° 28' E. 
Luting Shan: see Lutingkiao 

Ma-Chang-Kai, Tengyueh, Yunnan, China: 24° 39' N., 98° 00' E. 
Madaya Forest, Burma: 22° 30' N., 96° 06' E, 
Madischung Valley, Sikang, China: 26° 12' N., 101° 59' E. 
Mae Hong Song: see Changwat Mae Hong Son 
Mae Klong, Thailand: 13° 26' N., 100° 01' E. 

Mae Nam Khwae Noi, Kanchanaburi, Thailand: 14° 00' N., 99° 30' E. 
Mae Nam Yom, Prae, Thailand: 15° 52' N., 100° 16' E. 


Mae Pan, Phrae, Thailand: 18° 00' N., 100° 00' E. 

Mae Taw Forest, Thailand: 17° 20' N., 99° 35' E. 

Mae Tha Khwae, Tak, Thailand: 16° 54' N., 98° 52' E. 

Mae Wan River near Doi Saket, Thailand: 18° 50' E., 99° 35' E. 

Ma Fu Ling, Hupeh, China: 31° 52' N., 110° 18' E. 

Maha Oya, Ceylon: 07° 30' N., 81° 19' E. 

Ma Kai Hsien: see Makai 

Makai, Yunnan, China: 25° 42' N., 101° 55' E. 

Malipa, Burma: 23° 40' N., 98° 45' E. 

Maliwun, Burma: 10° 12' N., 98° 40' E. 

Mamoh: see Pakchan 

Mandalay, Burma: 21° 59' N., 96° 08' E. 

Mangpu, Bengal, India: 26° 58' N., 88° 27' E. 

M. Angton: see Muang Phrom Buri 

Mankeni, Ceylon: 08° 00' N., 81° 30' E. 

Manoram: see Ban Khung Samphao 

Mansum, Burma: 25° 47' N., 96° 15' E. 

Manthe, Burma: 25° 18' N., 95° 13' E. 

Maprau Island: see Ko Maphrao 

Margherita, Assam, India: 27° 17' N., 95° 47' E. 

Martaban, Burma: 16° 30' N., 97° 28' E. 

Matanga River, N. Kamrup, Assam: circa 26° 30' N., 91° 30' E. 

Matinga: see Matanga River 

Matsatap, Burma: 27° 30' N., 97° 50' E. ■ 

Matugama, Ceylon: 06° 30' N., 80° 08' E. 

Maungkan, Burma: 25° 05' N., 95° 00' E. 

Mawlaik, Burma: 23° 38' N., 94° 22' E. 

Mawlyngkueng, Assam, India: 25° 34' N., 92° 03' E. 

Mawphlang, Assam, India: 25° 27' N., 91° 46' E. 

Mawryngkueng: see Mawlyngkueng 

Maymyo, Burma: 22° 05' N., 96° 30' E. 

M. Chum Pon: see Ban Tha San 

Mee Nan, 30 miles N.E. of Uttaradit, Thailand: 17° 50' N., 100° 15' E. 

Mee Pan: see Mae Pan 

Mee Tan: see Ban Mae Tan Nua 

Mehongson: see Changwat Mae Hong Son 

Meiktila, Burma: 20° 50' N., 95° 52' E. 

Mekong-Salwin [Salween] Divide, Sikang, China: 28° 20' N., 99° 20' E. 

Mekong Valley, Yunnan, China: 28° 00' N., 99° 23' E. 

Mekong- Yangtze Divide, Yunnan, China: 27° 30' N., 99° 25' E. 

Melamoung: see Me Lamung 

Me Lamung, Thailand: 15° 32' N., 98° 48' E. 

Me Moh: see Ban Mae Mo 

Meng-ting, Yunnan, China: 23° 31' N., 99° 06' E. 

Me Ping River [Mae Ping], 180 miles N. of Bangkok, Thailand: circa 16° 40' N., 

99° 55' E. 
Mesarieng: see Ban Mae Sariang 
Me Taqua: see Mae Tha Khwae 
Me Taw Forest: see Mae Taw Forest 
Me Wong River, 53 mi. E. of Urn Pang, Thailand: 15° 50' N., 99° 38' E. 


Me Yam River: see Mae Nam Yom 

MiLi, Szechwan, China: 28° 11' N., 100° 50' E. 

MiNGUN, Sagaing, Burma: 22° 00' N., 95° 57' E. 

MiSHMi Hills, Assam, India: circa 28° 00' N., 96° 10' E. 

M. MouEN: see Moung Mouen 

MOKANSHAN, Chekiang, China: 30° 26' N., 119° 47' E. 

MOKLOK, Burma: 25° 37' N., 95° 25' E. 

MOKOKCHUNG, Assam, India: 26° 24' N., 94° 32' E. 

MONG Chit: see Ban Tuyong 

Mono Ha, N. Shan States, Burma: 22° 20' N., 98° 15' E. 

Mongkhor: see Ban Nong Kho 

MONGTZE, Yunnan, China: 25° 56' N., 112° 14' E. 

Moung Boum, Tonkin, Viet Nam: 22° 32' N., 102° 50' E. 

Moung Mo, Tonkin, Viet Nam: 22° 12' N., 102° 56' E. 

Moung Mouen, Tonkin, Viet Nam: 21° 30' N., 103° 00' E. 

Moung Moun, Tonkin, Viet Nam: 21° 36' N., 103° 20' E. 

Moung Yo, Laos: 21° 28' N., 101° 52' E. 

Mt. Angka: see Doi Inthanon 

Mt. Arizan, Formosa: 23° 32' N., 120° 46' E. 

Mt. Ava, Burma: 21° 52' N., 96° 01' E. 

Mt. Fan-si-pan, Tonkin, Viet Nam: 22° 19' N., 103° 49' E. 

Mt. Imaw Bum, Burma: 26° 08' N., 98° 30' E. 

Mt. Mooleyit: see Mulai-yit Hill 

Mt. Nwalabo, Burma: 14° 07' N., 98° 28' E. 

Mt. Omei, Szechwan, China: 29° 25' N., 103° 25' E. 

Mt. Popa, Burma: 20° 58' N., 95° 20' E. 

Mt. Tura, Assam, India: 25° 27' N., 90° 30' E. 

Mt. Victoria, Burma: 21° 17' N., 93° 53 'E. 

Mt. Wuchi: see Wuchi Shan 

Moupine: see Muping 

MUAKLEK [Muak Lek], Sraburi [Sara Buri], Thailand: 14° 35' N., 101° 17' E. 

Muang Chainat, Thailand: 15° 10' N., 100° 10' E. 

Muang Chiang Mai, Thailand: 18° 45' N., 99° 00' E. 

Muang Chiang Rai, Thailand: 19° 55' N., 99° 50' E. 

Muang Lampang, Thailand: 18° 18' N., 99° 31' E. 

Muang Nan, Thailand: 18° 47' N., 100° 47' E. 

Muang Nong Khai, Thailand: 17° 52' N., 102° 44' E. 

Muang Pai: see Ban Wiang Tai 

Muang Phichit: see Ban Nai Muang 

Muang Phitsanulok, Thailand: 16° 50' N., 100° 15' E. 

Muang Phrom Buri, Anghtong or Sing Buri, Thailand: 14° 50' N., 100° 26' E. 

Muang Prom: see Muang Phrom Buri 

Muang Qua Yie: see Nakhon Sawan 

Mu Cheng, Yunnan, China: 23° 45' N., 99° 10' E. 

Mudumalai, India: 11° 37' N., 76° 31' E. 

MuEK Lek: see Ban Muak Lek 

Mulai-yit Hill, Burma-Thailand border: 16° 05' N., 98° 43' E. 

MUNSIN, Burma: 25° 33' N., 95° 20' E. 

MuONG Boum, Tonkin, Viet Nam: 22° 32' N., 102° 50' E. 

MUONG Karbin: see Ban Kabin Buri 


MuONG Mo, Tonkin, Viet Nam: 22° 12' N., 102° 56' E. 

MuONG MoUN, Tonkin, Viet Nam: 21° 36' N., 103° 20' E. 

MuoNG Sen, Vinh, Viet Nam: 19° 32' N., 104° 07' E. 

MuONG Yo, Laos: 21° 28' N., 101° 52' E. 

Mupin: see Muping 

MUPING, Szechwan, China: 30° 29' N., 102° 49' E. 

Myawadi, Burma: 16° 41' N., 98° 31' E. 

Myitkyina, Burma: 25° 24' N., 97° 26' E. 

Nagarjun, Katmandu, Nepal: 27° 45' N., 85° 22' E. 

Nagchuka: see Hokow 

Naggenjung: see Nagarjun 

Na Hai, Tonkin, Viet Nam: 21° 12' N., 102° 55' E. 

Nakhon Fathom, Lum Phaya, Thailand: 13° 50' N., 100° 05' E. 

Nakhon Sawan, Thailand: 15° 41' N., 100° 07' E. 

Nakhon Si Thammarat, Thailand: 8° 26' N., 99° 58' E. 

Nakonrajasima [Nakhon Ratchasima], Thailand: 14° 57' N., 102° 05' E. 

Nakon Sawan: see Nakhon Sawan 

Nam Chuh: see Ban Nam Chut Yai 

Nam Fong, Hainan, China: 19° 24' N., 109° 33' E. 

Nam Khueng, Laos: 20° 25' N., 100° 18' E. 

Nam Nen, Tonkin, Viet Nam: circa 21° 40' N., 103° 15' E. 

Nam Tamai Valley, Burma: 27° 42' N., 97° 54' E. 

Nam Ting, Yunnan, China: 23° 31' N., 99° 06' E. 

Nam-ting River at Burma Border, China: 23° 23' N., 98° 43' E. 

Nan: see Muang Nan 

Nan-Tou Hsien, Wu Sheh, Taiwan, China: 23° 54' N., 120° 42' E. 

Nanyaseik, Burma: 25° 37' N., 96° 36' E. 

NAPfi, Laos: 18° 20' N., 105° 05' E. 

Narathiwat, Thailand: 6° 25' N., 101° 50' E. 

Na Tebbu land, Wantsang Ku, Kansu, China: circa 34° 00' N., 101° 30' E. 

Nauswa, Burma: 25° 40' N., 95° 20' E. 

Nawakot, Nepal: 27° 56' N., 85° 10' E. 

Nawocot: see Nawakot 

N'BUNGHKU, Burma: 25° 54' N., 96° 12' E. 

Nelliampathi Hills, India: 10° 30' N., 76° 35' E. 

Ngai Tio, Tonkin, Viet Nam: 22° 35' N., 103° 45' E. 

Ngan-son, Tonkin, Viet Nam: 22° 25' N., 106° 00' E. 

Ngapyiniun, Burma: 22° 32' N., 96° 00' E. 

Ngau-Tchi-Lea (Mts.), Hainan, China: 19° 00' N., 109° 30' E. 

Nghia Hung, Phu Qui, Annam: 19° 20' N., 105° 20' E. 

Ngoloshi: see Tsungshi 

Ngu-luko, Yunnan, China: 27° 03' N., 100° 12' E. 

Nhatrang, Annam, Viet Nam: 12° 15' N., 109° 15' E. 

NiKAWERATlYA, Ceylon: 07° 44' N., 80° 08' E. 

Nikawewa, near Kantalai, Ceylon: 08° 16' N., 81° 01' E. 

Nimrod Sound, Chekiang, China: 29° 40' N., 121° 50' E. 

NiNGPO, Chekiang, China: 29° 53' N., 121° 33' E. 

NiNGYUANFU, Szechwan, China: 27° 55' N., 102° 18' E. 

NiNGYUEN FU: See Ningyuanfu 

Ninh Hoa, Annam, Viet Nam: 12° 30' N., 109° 10' E. 


NODOA, Hainan, China: 19° 31' N., 109° 34' E. 

NOKREK, India: 25° 28' N., 90° 19' E. 

NoNG Bua: see Ban Nong Bua 

NoNG DOM Ta: see Ban Nong Don Ta 

NONGKAi: see Muang Nong Khai 

NONGKHOR: see Ban Nong Kho 

Nong Lum, Tonkin, Viet Nam: 22° 22' N., 102° 53' E. 

NONGPOH, Assam, India: 25° 55' N„ 91° 51' E. 

NONGSTOIN, Assam, India: 25° 31' N., 91° 16' E. 

Nong Yang, Thailand: 13° 10' N., 101° 30' E. 

NUA Chua Chan, Bien Hoa Prov., Viet Nam: 10° 56' N., 107° 23' E. 

Nui Kai, Yunnan, China: 26° 05' N., 100° 00' E. 

Nyetmaw River, Burma: 26° 05' N., 98° 18' E. 

Okma, Burma: 22° 35' N., 94° 56' E. 

Ok Pyam: see Pyam 

Ok Yam: see Pyam 

OOTACAMUND, Nilgiri Hills, India: 11° 24' N., 76° 44' E. 

Pa Ham, Tonkin, Viet Nam: circa 21° 35' N., 102° 55' E. 

Pahpoon: see Papun 

Pah Toop: see Pha Tup 

Pakchan: see Ban Pak Chan 

Pak Chong: see Ban Pak Chong 

Pakha: see Pa Kha 

Pa Kha, Tonkin, Viet Nam: 22° 33' N., 104° 18' E. 

Pak Hin Bun, Laos: 17° 35' N., 104° 40' E. 

Pak Jong: see Ban Pak Chong 

Pak Koh, Thailand: 18° 02' N., 99° 53' E. 

Paknampho: see Ban Pak Nam Pho 

PAK-Sfi, Laos: 15° 09' N., 105° 45' E. 

Paksong, Laos: 15° 15' N., 106° 07' E. 

Pak Tha, Thailand: circa 18° 00' N., 100° 00' E. 

Pak Tha Pujeg: see Ban Pak Tho 

Palasbari, Assam, India: 26° 10' N., 91° 35' E. 

Palghat, Malabar, India: 10° 40' N., 76° 35' E. 

Palni Hills, Madura Dist., Madras, India: circa 10° 20' N., 77° 30' E. 

Pang Mae Ton, Chiang Mai, Thailand: 18° 55' N., 99° 15' E. 

Pang Me Ton: see Pang Mae Ton 

Pang Nam Un, Nan, Thailand: circa 18° 30' N., 100° 33' E. 

Pang Sok: see Sathani Pang Sok 

Pantha, Burma: 23° 50' N., 94° 31' E. 

Paotaram: see Sathani Photharam 

Papun, Burma: 18° 05' N., 97° 22' E. 

Pasa, Laos: 19° 40' N., 102° 25' E. 

Pasbok, Darjeeling, Bengal, India: 27° 10' N., 88° 25' E. 

Pattipola, Ceylon: 06° 51' N., 80° 50' E. 

Pemionchee, Sikkim, India: 27° 20' N., 88° 17' E. 

Pennan (or Pangan) Island, Siam: see Ko Phangan 

Phanrang, Annam, Viet Nam: 11° 34' N., 108° 59' E. 

Pha Tup, Northern Thailand: South of Muang Nan 

Phawzaw, Burma: 25° 28' N., 95° 20' E. 


Phitsanuloke, Siam: 16° 50' N., 100° 15' E. 

Phong Saly, Laos: 21° 42' N., 102° 09' E. 

Phong Tho, Tonkin, Viet Nam: 22° 20' N., 103° 20' E. 

Phra Nakhon Si Ayutthaya, Ayutthaya, Thailand: 14° 20' N., 100° 35' E. 

Phu Do, Kan Luang, Nakon Phanom, Thailand: circa 16° 55' N., 104° 30' E. 

Phuket Island: see Ko Phuket 

Phu Kho, Kan Luang, Nakon Phanon, Thailand: circa 16° 55' N., 104° 30' E. 

Phu Kobo, Laos: 19° 22' N., 103° 23' E. 

Phu Phan, Sakon Nakon, Thailand: 16° 52' N., 104° 23' E. 

Phu Qui, Annam, Viet Nam: 19° 25' N., 105° 22' E. 

Piam: see DeLisle 

Picket: see Ban Nai Muang 

PiBN Ngai, Szechwan, China: 29° 12' N., 108° 10' E. 

Ping Tung, Taiwan, China: 22° 40' N., 120° 30' E. 

PiNTENNE, Ceylon: 07° 21' N., 81° 00' E. 

Pitsanulok: see Muang Phitsanulok 

Plateau Bolovens, Laos: 15° 10' N., 106° 20' E. 

Poi, Assam, India: 25° 20' N., 94° 37' E. 

PONMUDI, Travancore, India: 08° 44' N., 77° 07' E. 

POOKEIO: see Ban Phu Kheio 

Pookeiochaiyabhoom: see Ban Phu Kheio 

PoOTOO: see Puto Shan 

Pot ARAM: see Sathani Photharam 

PoWAi : see Poi 

Pra (Mt.), Cambodia: 12° 38' N., 103° 02' E. 

Prachuap Khiri Khan, Thailand: 11° 50' N., 99° 50' E. 

Pran: see Prachuap Khiri Khan and Pran Buri 

Pran Buri, Prachuap Khiri Khan, Thailand: 12° 22' N., 99° 56' E, 

Prapoot Mt.: see Khao Phra Phut 

Prapootabat: see Khao Phra Phut 

Prome, Burma: 18° 50' N., 95° 14' E. 

Pua: see Ban Pua 

Pucheng, Fukien, China: 28° 05' N., 118° 28' E. 

Pujeg: see Pujek 

PuJEK, Pak Tho, Rajburi, Thailand: 13° 23' N., 99° 42' E. 

Pukpitiya : see Puwakpitiya 

PULO Beman: see Clara Island 

PULO Ketchen: see St. Lukes Island 

PULO Lambi: see Sullivan's Island 

PULO Salakring: see Hastings Island 

PUM-KATONG, Burma: 25° 23' N., 97° 44' E. 

PUMSIN, Burma: 25° 57' N., 96° 11' E. 

PuTO Shan, Chusan Archipelago, Chekiang, China: 30° 00' N., 122° 25' E. 

Puwakpitiya, Gammaduwa, Ceylon: 07° 36' N., 80° 43' E. 

Pyam, Kampot, Thailand: 11° 37' N., 102° 56' E. 

Pyaungbyin, Burma: 24° 15' N., 94° 47' E. 

Pyaunggaung, Burma: 22° 35' N., 97° 05' E. 

Pyepat, Burma: 25° 54' N., 98° 15' E. 

Pynursla, Assam, India: 25° 18' N., 91° 53' E. 

Quangtri Phuoc Mon, Annam, Viet Nam: 16° 35' N., 107° 10' E. 


QUANG Tri River, Annam, Viet Nam: circa 16° 46' N., 107° 11' E. 

Racu Racu Mountains: see Rakuraku 

Raheng: see Ban Rahaeng 

Rahnampo: see Ban Pak Nam Pho 

Rajaburi: see Rat Buri 

Rajburi: see Ban Pong 

Rakuraku, Taiwan, China: 23° 27' N., 121° 30' E. 

Rama la Pass, Szechwan, China: 29° 40' N., 100° 25' E. 

Rangoon, Burma: 16° 47' N., 96° 10' E. 

Ranna, Ceylon: 06° 05' N., 80° 55' E. 

Rat Buri, Thailand: 13° 32' N., 99° 49' E. 

Ratnamti, Burma: 27° 35' N., 97° 55' E. 

Red Point, Tenasserim, Burma: 10° 40' N., 98° 30' E. 

Ringin, Sikkim, India: 27° 30' N., 88° 30' E. 

RiRi Bazaar, Nepal, India: 27° 55' N., 83° 22' E. 

Rumitchangu, Szechwan, China: 30° 50' N., 102° 05' E. 

RUNGBONG Valley, Sikkim, India: 26° 50' N., 88° 13' E. 

Sadiya, Assam, India: 27° 50' N., 95° 45' E. 

St. Lukes Island, Burma: 10° 05' N., 98° 15' E. 

Sakeo: see Ban Sa Kaeo 

Sakerat : see Ban Chakkrarat 

Salanga: see Koh Phuket 

Salangka: see Koh Phuket 

Salween-Mekong Divide at 28° 20' N.: 28° 20' N., 98° 35' E. 

Samasgi, Dharwar-Kanara border, India: 14° 40' N., 75° 00' E. 

Sambor, Cambodia: 12° 45' N., 105° 59' E. 

Sam roi Gop: see Sathani Sam Roi Yot 

Sangau, Assam, India: 21° 44' N., 93° 03' E. 

Sangsir: see Singsir 

Sata Hip: see Ban Sattahip 

Sathani Hin Lap, Thailand: 14° 40' N., 101° 10' E. 

Sathani Lamphura, Thailand: 07° 40' N., 99° 35' E. 

Sathani Pang Hua Phone, Thailand: 18° 25' N., 99° 15' E. 

Sathani Pang Sok, Sara Buri, Thailand: 14° 40' N., 101° 20' E. 

Sathani Photharam, Rat Buri, Thailand: 13° 40' N., 99° 50' E. 

Sathani Sam Roi Yot, Thailand: 12° 15' N., 99° 55' E. 

Sathani Tha Chomphu, Lamphun, Thailand: 18° 30' N., 99° 15' E. 

Sathani Thung Song, Nakhon Si Thammarat, Thailand: 08° 10' N., 99° 40' E. 

Sattha Hill: see Satthar 

Satthar, Nepal: 28° 01' N., 84° 35' E. 

Sawankaloki: see Ban Wang Mai Khon 

Sawankhalok: see Ban Wang Mai Khon 

Seechol: see Ban Sichon 

Sedonchen, Sikkim, India: 27° 30' N., 88° 30' E. 

Se'en, N. Shan States, Burma: 22° 39' N., 97° 22' E. 

Seniku, Burma: 25° 34' N., 97° 47' E. 

Sevoke, Bengal Presidency, Pakistan: 26° 51' N., 88° 32' E. 

Shandaw, Burma: 17° 55' N., 95° 40' E. 

Shanghai, China: 31° 13' N., 121° 25' E. 

Shang Kuan, Yunnan, China: 25° 55' N., 100° 05' E. 


Shihpingchow, Yunnan, China: 23° 40' N., 102° 25' E. 

Shinei, Formosa, China: 23° 18' N., 120° 18' E. 

Shineigun: see Shinei 

Shingbwiyang, Hukawng Valley, Burma: 26° 41' N., 96° 12' E. 

Shoo-O-Lo: see Siolo 

Shrivilliputtur [Srivilliputtur], Madras, India: 07° 30' N., 77° 38' E. 

Shuan Lung Chang, Kweichow, China: 28° 05' N., 106° 48' E, 

Shui-Kow-Kwan: see Lungchow 

SHWEBO, Burma: 22° 35' N., 95° 42' E. 

Shweli Valley, Yunnan, China: circa 25° 00' N., 98° 40' E. 

Shweli-Salween Divide, Yunnan, China: circa 25° 05' N., 98° 45' E. 

SiAOMENGTUNG, Yunnan, China: 24° 10' N., 99° 15' E. 

SiEM Reap, Cambodia: 13° 20' N., 103° 51' E. 

Siken: see Ban Si Khiu 

Sikeu: see Ban Si Khiu 

SiMA, Myitkyina, Burma: 25° 05' N., 97° 35' E. 

Singkaling Hkamti, Burma: 26° 00' N., 95° 39' E. 

SiNGSiR, Bengal Presidency, India: 27° 04' N., 88° 35' E, 

Singtam, Sikkim, India: 27° 14' N., 88° 33' E. 

SiNKAi, Yunnan, China: 24° 12' N., 102° 04' E. 

Siolo, Szechwan, China: 30° 02' N., 100° 48' E. 

Siracha: see Ban Si Racha 

SiRSA, Hissar, Punjab, Pakistan: 29° 30' N., 75° 00' E. 

SiSAGARHi, Nepal: 27° 25' N., 85° 05' E. 

Slokbai: see Ban Salok Bat 

SoKOTAi: see Ban Thani 

SOOKIA POKHARi, Darjeeling, India: 27° 02' N., 88° 14' E. 

SOOKLI: see Sukli 

South Cape of Formosa, China: 21° 54' N., 120° 52' E. 

Sriracha : see Ban Si Racha 

SUCHOW, Szechwan- Yunnan border, China: 28° 43' N., 104° 32' E. 

SuiFU: see Suchow 

SuiYANG, Kweichow, China: 27° 50' N., 107° 18' E. 

SUKHOTHAi: see Ban Thani 

SUKIAPOKHRI : see Sookia Pokhari 

SUKLI, Myawadi Rd., Burma: 16° 42' N., 98° 22' E. 

Sullivan's Island, Mergui Archipelago, Burma: 10° 40' N., 97° 58' E. 

SUMKA Uma, Burma: 25° 57' N., 97° 49' E. 

SUMPRABUM, Burma: 26° 38' N., 97° 36' E. 

Sungei Balik, Tenasserim, Burma: 10° 31' N., 98° 33' E. 

Sungpan, Szechwan, China: 32° 41' N., 103° 21' E. 

Sunjang: see Suiyang 

Sung Shan, Honan, China: 34° 35' N., 113° 20' E. 

Ta Chang Tai: see Tha Chang Tai 

Ta Chiao, Szechwan, China: circa 30° 03' N., 103° 13' E. 

Tachin: see Ban Maha Chai 

Taga Hka, Burma: 26° 22' N., 96° 07' E. 

Tagoot, Burma: 10° 25' N., 99° 18' E. 

Ta-ho: see Ta Ho-Hta 

Ta Ho-Hta, Independent Kareni, Burma: 19° 06' N., 97° 30' E. 


Tainan, Taiwan, China: 23° 00' N., 120° 12' E. 

Tai-Pa-Shiang Mts. : see Tapa shan 

Taipei, Taiwan, China: 25° 05' N., 121° 32' E. 

Taipei Hsien, Wu Lai, Taiwan, China: 24° 50' N., 121° 32' E. 

Tai-pei-shan District, Shensi, China: 33° 56' N., 107° 41' E. 

Tai-ping-pu, Yunnan, China: 24° 57' N., 98° 42' E. 

Tai-yuan-fu, Shansi, China: 37° 53' N., 112° 29' E. 

Tak: see Ban Rahaeng 

Takubama, Assam, India: 25° 50' N., 94° 28' E. 

Tamabin, Burma: circa 17° 45' N., 96° 30' E. 

Tamanthi, Burma: 25° 20' N., 95° 12' E. 

Tam Dao, Tonkin, Viet Nam: 21° 30' N., 105° 38' E. 

Tamu, Burma: 25° 45' N., 98° 02' E. 

Tanga, Burma: 25° 50' N., 98° 05' E, 

Tangshu, Gieu Long Shien, Tibet: 28° 58' N., 98° 08' E. 

Tang Gu: see Tangshu? 

Tanjong Badak, Tenasserim, Burma: 10° 06' N., 98° 29' E, 

Taok, Tenasserim, Burma: 16° 20' N., 98° 28' E. 

Tag Mung Chung, Lu-tien, Yunnan, China: 27° 12' N., 103° 31' E. 

Tapa Hka, Burma: 26° 07' N., 96° 15' E. 

Tapa shan, Shensi, China: 32° 40' N., 107° 30' E. 

Tappan-sha, Formosa, China: 23° 28' N., 120° 44' E. 

Tapposha: see Tappan-sha 

Tarkhola, Sikkim, India: 27° 07' N., 88° 33' E. 

Taroar: see Tagoot 

Taron Valley, Burma: circa 28° 00' N., 98° 10' E. 

Ta-shui-tang: see Tashutang 

Tashutang, Yunnan, China: 24° 17' N., 99° 15' E. 

Tasu Bum, Burma: 26° 04' N., 96° 14' E. 

Tatkon, Burma: 23° 45' N., 94° 23' E. 

Tatsienlu, Szechwan, China: 30° 03' N., 102° 13' E. 

Taukchandmai, Burma: 23° 30' N., 94° 35' E. 

Tavoy, Burma: 14° 07' N., 98° 18' E. 

Tavoy Island, Mergui Archipelago, Burma: 11° 00' N., 98° 20' E. 

Tawmaw, Burma: 25° 44' N., 96° 18' E. 

Tay Ninh, Cochin China, Viet Nam: 11° 12' N., 106° 02' E 

Tehan, China: circa 30° 20' N., 117° 20' E. 

Tellula, Ceylon: 06° 36' N., 81° 08' E. 

Telok Besar, Tenasserim, Burma: 10° 22' N., 98° 35' E. 

Tempao, Burma: 24° 59' N., 94° 56' E. 

Tenasserim Town, Burma: 12° 06' N., 99° 03' E. 

Tengyueh, Yunnan, China: 25° 00' N., 98° 30' E. 

Teraso : see Teraso Soan 

Teraso Soan, Formosa, China: 22° 01' N., 120° 51' E. 

Tha Chang: see Ban Tha Chang 

Tha Chang Tai, Tak, Thailand: 16° 51' N., 99° 03' E. 

Thagyet, Burma: 12° 06' N., 99° 06' E. 

Thai Nguyen, Tonkin, Viet Nam: 21° 30' N., 105° 55' E. 

Thai Nien: see Thai Nguyen 

Thale Song Hong, Thailand: 7° 50' N., 99° 27' E. 


Thamaung: see Thandaung 

Thandaung, Burma: 19° 05' N., 96° 38' E. 

Than Hoa, Annam, Viet Nam: 19° 49' N., 105° 48' E. 

Tha Rong: see Ban Wichian 

Thateng: see Ban Thateng 

Thaton, Burma: 16° 50' N., 97° 18' E. 

Thatone: see Thaton 

Thaungyin Valley, Tenasserim, Burma: circa 16° 42' N., 98° 30' E. 

Theng-gan-ngok, Tenasserim, Burma: 13° 20' N., 98° 55' E. 

Thobal, Manipur, India: 24° 18' N., 94° 02' E. 

Thotel: see Thobal 

Thowngyoh: see Theng-gan-ngok 

Thua Luu, Annam, Viet Nam: circa 16° 19' N., 108° 00' E. 

Thuangyin Valley: see Thaungyin Valley 

TiENCHUAN, Szechwan, China: 30° 05' N., 103° 00' E, 

TiNGCHOW, Hopeh, China: 38° 30' N., 115° 02' E. 

Tingchowfu, Fukien, China: 25° 44' N., 116° 27' E. 

TiNNPANi: see Tirappane 

TiRAPPANE, Ceylon: 08° 12' N., 80° 31' E. 

Ti-yu Gomba, Yunnan [Sikang], China: circa 28° 40' N., 101° 10' E. 

Tongka: see Koh Phuket 

TOONG, Sikkim, India: 27° 35' N., 88° 40' E. 

Torsan: see Ban Tha San 

TOUNGOO, Burma: 18° 55' N., 96° 25' E. 

Tra Khanun: see Ban Tha Khanum 

Trang Bom, Cochin China, Viet Nam: 10° 50' N., 107° 00' E. 

Trashi-cho-ten, Szechwan, China: circa 30° 20' N., 102° 20' E, 

Trivandrum, Travancore, India: 08° 29' N„ 76° 55' E. 

TsAO Po, Szechwan, China: 21° 05' N., 103° 31' E. 

TSE-KOW: see Tseku 

TsEKU, Yunnan, China: 28° 00' N., 98° 52' E. 

TsEO JiA Keo, Szechwan, China: circa 28° 18' N., 104° 12' E. 

TsiNGKiEN, Shensi, China: 37° 12' N., 110° 12' E. 

TsiNG-LiNG Mts., Shensi, China: circa 33° 00' N., 108° 00' E. 

TsiNGTEH, Anhwei, China: 30° 23' N., 118° 30' E. 

TSONMA, Burma: 26° 10' N., 98° 35' E. 

TsuK KON Shih: see Tungshih 

TsUNGSHi: see Tungshih 

TSUNYI HsiEN, Kweichow, China: 27° 35' N., 107° 02' E. 

TUNG-LiNG: see Eastern Tombs 

TUNGLU, Chekiang, China: 29° 47' N., 119° 37' E. 

Tung Nog Karien, Jombing, Rajburi, Thailand: 13° 33' N., 99° 35' E. 

Tung Sawn: see Sathani Thung Song 

Tungshih, Hupeh, China: 30° 22' N., 111° 42' E. 

TUNGTZE, Kweichow, China: 27° 58' N., 106° 51' E. 

Tung Wong Tien, Kweichow, China: 28° 03' N., 105° 50' E. 

Tu-PA-KEO, Moupine, Szechwan, China: circa 30° 30' N., 102° 45' E. 

Tura, Garo Hills, Assam, India: 25°31'N., 90°16'E. 

Ukhrul, Manipur, India: 25° 10' N., 94° 18' E. 

Ulongkong, Szechwan, China: circa 29° 55' N., 102° 10' E. 


Um Pang: see Ban Le Kathe 

Umran, Assam, India: 25° 65' N., 91° 50' E. 

Uttaradit, Thailand: 17° 38' N., 100° 06' E. 

Van Lung, Thanhoa, Annam, Viet Nam: 20° 30' N., 104° 40' E. 

ViCHiANBURi: see Ban Wichian 

Victoria Point, Tenasserim, Burma: 10° 00' N., 98° 30' E. 

Vientiane, Laos: 17° 45' N., 102° 40' E. 

Wan Mts., Anwei, China: circa 30° 45' N., 117° 55' E. 

Wang Kien: see Kanchanaburi 

Wan hsien, Szechwan, China: 30° 49' N., 108° 23' E. 

Wan-tien, Yunnan, China: 24° 31' N., 99° 21' E. 

Wa Shan, Szechwan, China: circa 29° 25' N., 104° 20' E. 

Waterfall, Mekang: see Ban Mae Klang 

Wa TIEN, China: 25° 20' N., 98° 36' E. 

Wei Chow: see Weikiu 

Weikiu, Szechwan, China: 31° 25' N., 103° 28' E. 

Weisi Pass, Yunnan, China: 27° 14' N., 99° 27' E. 

Wellawaya, Uva Prov., Ceylon: 06° 43' N., 81° 06' E. 

Wenchwan, Szechwan, China: 31° 20' N., 103° 31' E. 

Wen-shui, Kweichow, China: 28° 25' N., 106° 18' E. 

Western Hills, 15 mi. W. of Peking, Hopeh, China: 39° 57' N., 116° 07' E. 

WiE Shi Pass: see Weisi Pass 

Wu-chi: see Wushi 

WuCHi Shan, Hainan, China: 18° 55' N., 109° 20' E. 

WUNG Pratart Farm, Kam Peng Pet, Thailand: 16° 10' N., 99° 45' E. 

WUNTHO, Burma: 23° 56' N., 95° 45' E. 

WuSHi, Yunnan, China: 29° 05' N., 101° 28' E. 

WUTAI, Shansi, China: 38° 55' N., 113° 38' E. 

WuTiNGCHOW, Yunnan, China: 25° 30' N., 102° 26' E. 

WuTiNG Hsien: see Wutingchow 

Wu-TING-SHAN, Jehol, Manchuria: 40° 42' N., 117° 21' E. 

Wu-TSAi: see Wutai 

XiENG Khouang, Laos: 19° 25' N., 103° 25' E. 

XiEN QUANG Koo: see Xieng Khouang 

Yachow: see Yachowfu 

Yachowfu, Szechwan, China: 29° 59' N., 103° 10' E. 

Yalung River, S.W. Szechwan, China: circa 27° 10' N., 101° 50' E. 

Yangwupa, Yunnan, China: 23° 56' N., 102° 10' E. 

Yao-shan, Kwangsi, China: 24° 00' N., 110° 00' E. 

Yellapur, Bombay, India: 14° 55' N., 74° 40' E. 

Yen-an-fu, Shensi, China: 36° 44' N., 109° 25' E. 

Yenping, Fukien, China: 26° 38' N., 118° 10' E. 

Yin, Burma: 22° 43' N., 94° 42' E. 

Yiu Shan: see Yu Shan 

YUKi, Fukien, China: 25° 28' N., 119° 18' E. 

Yulong-Vlang: see Ulongkong 

Yung Cha Shan, Szechwan, China: 29° 23' N., 108° 10' E. 

Yunghtang, Burma: 27° 38' N., 98° 03' E. 

Yung-Ning, Yunnan, China: 27° 45' N., 100° 41' E. 

Yungpeh, Szechwan, China: 26° 40' N., 100° 45' E. 


Yung-pei-ting: see Yungpeh 

YUNNANFU, Yunnan, China: 25° 00' N., 102° 40' E. 

YUNNANYi, Yunnan, China: 25° 25' N., 100° 37' E. 

Yu SHAN, Kwangtung, China: 24°42'N.,114°10'E. 

Zamayi Res., Pegu Yoma, Burma: 18° 21' N., 95° 59' E. 

Zami River, 100 miles south of Moulmein, Burma: 15° 25' N., 98° 25' E. 

Zaulep Ga, Burma: circa 25° 57' N., 96° 11' E. 

Zaungtu, Burma: circa 17° 50' N., 96° 30' E. 


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1914. Mammals collected or observed by the Swedish zoological expedition to 
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1936. The penis bone in the Ceylonese squirrels, with special reference to its 
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1949. Notes on the snakes and mammals of the High Wavy Mountains, Madura 
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1883. List of the specimens of squirrels in the Leyden Museum. Notes from 
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1936. The Vernay-Hopwood Upper Chindwin expedition. Jour. Bombay Nat. 
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1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic expeditions. 
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Wang, Chi-Wu 

1956. China, vegetation (description and map). Encyclopedia Britannica 1956 
Ed., 5, p. 513. 

Wroughton, R. C. 

1905. "The" common striped palm squirrel. Jour. Bombay Nat. Hist. Soc, 
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1908. On the forms of squirrel hitherto classed under Sc. finlaysoni Horsf . Ann. 
Mag. Nat. Hist. (ser. 8), 2, pp. 393-401. 

1910. Some notes on the giant squirrels of India, Burma, and Ceylon. Jour. 
Bombay Nat. Hist. Soc, 19, no. 4, pp. 880-896. 

1912. The Bombay Natural History Society's mammal survey of India, Jour. 
Bombay Nat. Hist. Soc, 21, no. 4, pp. 1170-1196. 

1915a. Bombay Natural History Society's mammal survey of India, Burma 
and Ceylon. Report No. 16; Dry zone, central Burma, and Mt. Popa. Jour. 
Bombay Nat. Hist. Soc, 23, pp. 460-480. 

1915b. Bombay Natural History Society's mammal survey of India, Burma 
and Ceylon. Report No. 17: South Tenasserim. Jour. Bombay Nat. Hist. 
Soc, 23, pp. 695-720. 

1915c. Bombay Natural History Society's mammal survey of India, Burma 
and Ceylon. Report No. 19: Bengal, Bihar and Orissa. Jour. Bombay Nat. 
Hist. Soc, 24, pp. 96-110. 

1916a. Bombay Natural History Society's mammal survey of India, Burma 
and Ceylon. Report No. 23: Sikkim and Bengal Terai. Jour. Bombay Nat. 
Hist. Soc, 24, pp. 468-493, map. 

1916b. Bombay Natural History Society's mammal survey of India, Burma 
and Ceylon. Report No. 20: Chindwin River. Jour. Bombay Nat. Hist. 
Soc, 24, pp. 291-309. 

1916c Scientific results from the mammal survey. No. 13. G. New rodents 
from Sikkim. Jour. Bombay Nat. Hist. Soc, 24, pp. 424-430. 

1916d. Scientific results from the mammal survey. No. 14. D. The squirrels 
of the Funamhulus palmarum-tristriatus group in the peninsula. Jour. Bom- 
bay Nat. Hist. Soc, 24, pp. 644-649. 

1919. Summary of the results from the Indian Mammal Survey of the Bombay 
Natural History Society. Part III. Jour. Bombay Nat. Hist. Soc, 26, no. 2, 
pp. 338-379. 

1921, On the erythraeus group of squirrels. Jour. Bombay Nat. Hist. Soc, 27, 

pp. ns-m. 

Wroughton, R. C, and W. M. Davidson 

1919. Scientific results from the mammal survey. No. 20. C. Two new forms 
of the "Funambulus tristriatus" group. Jour. Bombay Nat. Hist. Soc, 26. 
pp. 728-730. 

Zahn, Walter 

1942. Die Riesen-, Streifen-, und Spitsnasenhornchen der Orientalischen Re- 
gion. Zeitschrift fur Saugetierkunde, 16, no. 1, pp. 1-182, 6 maps. 


Abdulali, Humayun 9, 44, 47 

= Dremomys rufigenis rufigenis 280 

Callosciums caniceps 189 
affinis, Ratufa 33 
agouti pelage 14 

= Callosciurtis flavimanus styani 152 

Callosciurus finlaysoni 167 
Allen. G. M. (1940) 20 

Callosciurus caniceps 195 

= Menetes berdmorei berdmorei 297 
Anderson, John (1878) 18 

=Funambulus tristriatus tristriatxis 87 

Callosciurus finlaysoni 172 
Anthony, Harold E. 9 

— Callosciurus erythraetis intermedius 110 
Archbold Expeditions 7 

Funambulus pennanti 76 

= Callosciurus pygerythrus lokroides 227 

Callosciurus flavimanus 138 


= Callosciurus 99 

Tamiops mcclellandi 241 

Callosciurus erythraeus 117 

= Callosciurus flavimanus quinqu£striatus 

Dremomys rufigenis 282 

Funambulus palmarum 81 

== Callosciurus pygerythrus m^arsi 222 

Funambulus palmarum 82 

= Ratufa indica maxima 45 

= Callosciurus caniceps domelicus 195 

berdmorei, Menetes 294 

Menetes berdmorei 297 

= Dremomys lokriah lokriah 264 

Callosciurus erythraeus 108 
bicolor, Ratufa 49 

Callosciurus caniceps 190 
Biswas, Biswamoy 9 
Black, Craig C. 27 
Blanford (1888-91) 19 

= Callosciurus phayrei 201 

Callosciurus pygerythrus 226 

Callosciurus finlaysoni 179 

= Callosciurus flavimanus flavimanus 146 

= Ratufa indica indica 43 

Callosciurus flavimanus 151 

Callosciurus finlaysoni 184 

Funambulus palmarum 85 
Burt, W. H. 8 


Dremomys pernyi 276 
Callosciurus 99 

caniceps 187 

erythraeus 101 

ferrugineus 158 

flavimanus 120, 189 

finlaysoni 161 

inornatus 209 

phayrei 201 

pygerythrus 213 
caniceps, Callosciurus 187 

Callosciurus caniceps 196 

= Callosciurus flavimanus styani 152 

= Callosciurus erythraeus bartoni 117 
Carter, T. Donald 305 

Callosciurus caniceps 194 





Callosdurus flavimanus 143 

= Ratufa macroura macroura 37 

=Ratufa bicolor phaeopepla 52 

Ratufa indica 47 

= Ratufa macroura macroura 37 

=Dremomys pernyi calidior 276 

= Callosdurus caniceps caniceps 196 

Callosdurus finlaysoni 170 

= Tamiops swinhod swinhod 248 
classification 20 

= Callosdurus finlaysoni bocourti 179 

—Sdurotamias davidianus consobrinus 

Tamiops m^clellandi 239 
color names 14 

Funximhulus palmar um 81 

Callosdurus caniceps 187 

Ratufa bicolor 58 

Sdurotamias davidianus 307 

Menetes berdmord 300 

= Callosdurus flavimanus flavimanus 146 
Cranbrook, Lord 63, 113, 120, 121, 239, 250, 


=Callosdurus erythraeus erythrogaster 105 

= Callosdurus erythraeus bhutanensis 108 


= Callosdurus flavimanus flavimanus 146 

Ratufa macroura 38 
davidianus, Sdurotamias 304 

Sdurotamias davidianus 307 
Davis, D. Dwight 291 

= Callosdurus caniceps bimacuJatus 190 

Ratufa indica 41 

Menetes berdmord 299 
Deignan, Herbert G. 9 


=Funambulus sublineatus sublineatus 95 
Deraniyagala, P. E. P. 9 

= Callosdurus finlaysoni dnistralis 177 
dimensions 15 
diurnal squirrels 10, 12 

= Tamiops rodolphd rodolphd 244 

Callosdurus caniceps 195 
Dorst, M. Jean 8 

Funambulus layardi 94 
Dremamys 11, 259 

everetti 17, 289 

lokriah 263 

pernyi 269 

pyrrhomerus 283 

rufigenus 278 

= Funambulus tristriatus tristriatus 87 


Tamiops rodolphd 245 
EUerman (1940) 20 

—Ratufa indica indica 43 

= Ratufa 29 

= Callosdurus caniceps bimaculatus 190 

= Callosdurus caniceps concolor 187 
erythraeus, Callosdurus 101 

Callosdurus erythraeus 104 

Callosdurus erythraeus 105 

= Callosdurus 99 
everetti 11 

everetti, Dremomys 17, 289 
evolution 7, 15 
evolutionary implications 8 
extraterritorial 11 


Callosdurus caniceps 195 

Funambulus palmarum 83 

Ratufa bicolor 58 
ferrugineus, Callosdurus 158 
finlaysoni, Callosdurus 161 

Callosdurus finlaysoni 165 
flavimanus, Callosdurus 120 

Callosdurus flavimanus 146 

Dremomys pernyi 275 




= Callosciurus finlaysoni bocourti 179 

= Callosciurus caniceps caniceps 196 

Callosciurus finlaysoni 166 
Forcart, L. 8 

= Tamiops maritimus maritimus 252 
forresti, Sciurotamias (Rupestes) 309 

= Tamiops sunnhoei sunnhoei 248 

Callosciurus finlaysoni 167 

— Callosciurus erythraeus bartoni 117 

= Callosciurus flavimanus griseimanus 148 
Funambulus 65 

layardi 91 

palmarum 77 

pennanti 72 

sublineatus 95 

tristriatus 87 

= Dremomys rufigenis rufigenis 280 


Dremomys lokriah 266 

Callosciurus finlaysoni 169 

Ratufa bicolor 59 

Callosciurus flavimanus 150 

Callosciurus flavimanus 129 

= Funambulus palmarum palmarum 79 
Granger, Walter 287 
Gray (1867) 18 

Callosciurus flavimanus 148 

= Dremomys pernyi pernyi 271 
griseopectus Blyth, 1847 

= Callosciurus flavimanus shanicus 131 
griseopectus Thomas, 1894 

= Callosciurus flavimanus styani 152 

= Callosciurus finlaysoni sinistralis 177 
Dremomys pyrrhom^rus 288 

= Callosciurus flavimanus michianus 134 

Ratufa bicolor 63 

Tamiops maritimus 255 

Callosciurus erythraeus 118 


Callosciurus finlaysoni 167 

= Callosciurus caniceps bimaculatus 191 
Hayman, R. W. 8 

= Callosciurus phayrei 201 

= Callosciurus caniceps caniceps 196 

= Callosciurus caniceps caniceps 196 

Callosciurus flavimanus 143 

= Callosciurus finlaysoni cinnamomeus 170 
Hershkovitz, Philip 8 

= Callosciurus 99 
Holm, A. 9 

= Tamiops rodolphei rodolphei 244 
Hooper, Emmett T. 8 
Hora, Sunder Lai 9 
Horsfield (1824) 18 

Dremomys pernyi 273 
Husson, A. M. 9 
Hyosciurus 10 

Callosciurus flavimanus 133 


= Callosciurus flavimantis quinquestriatus 

= Callosciurus inornatus 209 

= Dremomys pernyi howelli 273 

Tamiops m/;clellandi 240 
Indian Subregion 10, 12 
indica, Ratufa 41 

Ratufa indica 43 

= Funambulus palmarum palmarum 79 
Indochinese Subregion 10, 12 

= Callosciurus caniceps caniceps 196 
inornatus, Callosciurus 209 

= Callosciurus flavimanus castaneoventris 

Callosciurus erythraeus 110 

Callosciurus pygerythrus 218 
Jentink (1883) 18 
Johnson, David H. 8 




= Funambulus sublineatus obscurtis 97 

Funambulus palmarum 84 

-Callosciurus erythraeus sladeni 113 

= Callosciurus ferrugineus 158 

= Callosciurus erythraeus erythrogaster 105 

Tamiops mcclellandi 240 
Koopman, Karl F. 8 

=Menetes berdmorei mouhotei 301 


=Dremomys rufigenis rufigenis 280 

= Tamiops mxiritimus liainanus 255 

^Callosciurus caniceps concolor 187 

= Sciurotamias davidianus datridianus 30 
Lawrence, Barbara 8 
layardi, Funambulus 93 

Funambulus layardi 93 

=Dremomys pernyi pernyi 271 

= Callosciurus finlaysoni bocourti 179 

= Callosciurus finlaysoni bocourti 179 

Ratufa bicolor 55 

= Callosciurus flavim^imis griseimxinus 148 

Tamiops mcclellandi 242 

= Callosciurus flavimanus styani 152 

= Tamiops rodolphei rodolphei 244 

= Dremomys pernyi pernyi 271 
locality 11 

lokriah, Dremomys 263 

Dremomys lokriah 264 

Callosciurus pygerythrus 227 

= Callosciurus caniceps bimaculatus 190 

Funambulus pennanti 75 

= Ratufa bicolor gigantea 59 

= Callosciurus finlaysoni sinistralis 177 

= Tamiops rodolphei rodolphei 244 


Dremomys lokriah 268 
mxicroura, Ratufa 34 

Ratufa Ttiacroura 37 

= Ratufa bicolor gigantea 59 

= Ratufa macroura macroura 37 
Major (1893) 19 

= Ratufa indica maxima 45 
Malaysian Subregion 12 

= Tamiops mcclellandi mcclellandi 237 

= Callosciurus caniceps bimaculatus 190 

= Ratufa bicolor phaeopepla 52 
maritimus, Tamiops 251 

Tamiops maritimus 252 

= Callosciurus caniceps bimxiculatus 190 

= Funambulus palmarum favonicus 83 

Ratufa indica 45 
McGann, Charles 70, 73 
mcclellandi, Tamiops 235 

Tamiops mcclellandi 237 

Callosciurus pygerythrus 222 
Meise, W. 9 

Ratufa macroura 38 

= Dremomys pyrrhomerus riudonensis 288 

Callosciurus finlaysoni 176 
Menetes 293 

berdmorei 294 

= Dremomys pernyi howelli 273 
Mertens, R. 9 

Callosciurus flavimanus 134 

= Callosciurus erythraeus sladeni 113 

= Callosciurus erythraeus bartoni 117 

= Callosciurus caniceps bimaculatus 190 

= Dremomys pernyi senex 276 

Menetes berdmorei 299 

= Callosciurus caniceps adangensis 189 
= Callosciurus caniceps adangensis 189 




Tamiops maritimtis 256 

=Ratufa macroura macroura 37 

Tamiops m^ritimus 253 
Morrison-Scott, T. C. S. 8 

Menetes herdmorei 301 


= Callosciurus erythraeus erythrogaster 105 

= Callosciurus caniceps bimaculatus 190 
National Science Foundation 9 

Callosciurus flavimanus 154 

= Tamiops m^clellandi leucotis 242 

Callosciurus finlaysoni 169 

Funambulus tristriattis 91 


Funambulus sublineatu^ 97 

= Tamiops sivinhoei swinhoei 248 

Funambulus palmarum 86 

Dremomys rufigenis 283 
order 16 

Oriental Region 10 
original description 17 

= Dremomys rufigenis rufigenis 280 
Osgood (1934) 20 

Callosciurus pygerythrus 220 

Dremxymys pernyi 277 

=Sciurotamias davidianus consobrinus 308 


Drem,omys lokriah 267 
palynarum, Funambulus 77 

Funambulus palmxirum 79 

= Funambulus 65 

= Callosciurus caniceps bimaculatus 191 

^Callosciurus caniceps bimaculatus 190 
pelage color 17 

= Tamiops mcclellandi m^clellandi 237 

= Funambulus palmarum palmarum 79 


Menetes berdmorei 299 
pennanti, Funambulus 74 

Funambulus pennanti 74 
pernyi, Dremomys 269 

Dremomys pernyi 271 
Petauristinae 10 

Ratufa bicolor 52 

= Callosiurus flavimanus griseimanus 184 

= Callosciurus caniceps bimaculatus 191 

= Callosciurus flxivimanus flavimanus 146 
Pocock (1923) 19 

= Callosciurus finlaysoni finlaysoni 165 

= Callosiurus finlaysoni bocourti 179 

Callosiurvs flavimanus 141 
Prasad, M. R. N. 72 
Prasadsciurus 71 

= Callosciurus flavimanus zimmeensis 136 
procedure 7, 8, 11, 13-17 
Prosciurillus 10 
Public Health Service 9 

= Callosciurus erythraeus erythrogaster 105 

= Ratufa indica indica 43 
pygerythrus, Callosciurus 213 

Callosciurus pygerythrus 217 

= Menetes berdmorei mouhotei 301 
pyrrhomerus, Dremxrmys 283 

Dremomys pyrrhomerus 287 


= Callosciurus flavimanus flavimanus 146 

Callosciurus flavimanus 126 

= Callosciurus finlaysoni bocourti 179 
72a<M/a 29 
affinis 33 
bicolor 49, 11 
indica 41 
macroura 34 

Dremomys pyrrhomerus 288 

= Tamiops maritimus hainanus 255 
Roberts, H. Radclyffe 8 

Funambulus palmarum 82 



Robinson and Kloss (1918) 19 
rodolphei, Tamiops 243 

Tamiops rodolphei 244 
Rubrisdunis 10 

Callosciurus flavimanus 142 

= Callosciurus erythraeus sladeni 113 
rufigenis, Dremomys 278 

Dremomys rufigenis 280 

= Ratufa 29 
Rupestes 309 

forresti 309 

= Tamiops swinhoei swinkoei 248 


= Sciurotamias davidianus consobrinus 308 

= Callosciurus caniceps bimaculatus 190 
Sanborn, Colin C. 8 

= Tamiops maritimus m^riiimus 252 
Sdurotamias 10, 27, 303 

davidianus 304 

/orresfi 309 

Dremomys pernyi 276 

Callosciurus flavimanus 131 

= Callosciurus erythraeus bartoni 117 

Callosciurus flavimanus 140 

Funambulus layardi 93 

= Callosciurus pygerythrus lokroides 227 

= Ratufa macroura dandolena 38 

Callosciurus finlaysoni 177 

Ratufa bicolor 56 
skull character 16 
skull measurements 16 

Callosciurus erythraeus 113 
Smith, Floyd T. 9 

Ratufa bicolor 57 
Sody, H. J. V. 9 

= Callosciurus erythraeus haringtoni 118 

= Tamiops svrinhoei surinhoei 248 

= Callosciurus finlaysoni cinnamameus 170 


Callosciurus pygerythrus 224 

= Ratufa bicolor hainana 63 
Stop-Bowitz, C. 9 
Stresemann, E. 9 

Callosciurus flavimanus 152 

= Dremomys lokriah lokriah 264 
sublineatus, Funambulus 95 

Funambulus sublineatus 95 

= Callosciurus caniceps bimaculatus 190 

= Ratufa indica indica 43 
swinhoei, Tamiops 246 

Tamiops swinhoei 248 

= Callosciurus flavimanus quinquestriatus. 
synonymy 15 


= Callosciurus finlaysoni bocourti 179 

= Callosciurus flavimnnus siamensis 140 

= Callosciurus caniceps bimaculatus 191 
Tamiasciurini 27 
Tamiasciurus 27 

= Funambulus 65 
Tamiops 231 

maritimus 251 

mcclellandi 235 

rodolphei 243 

sunnhoei 246 
Tate, George H. H. 7 

= Callosciurus caniceps bimaculatus 191 
taxonomic history 18 

= Callosciurus caniceps concolor 187 

= Ratufa macroura macroura 37 

= Callosciurus caniceps adangensis 189 

Callosciurus flavimanus 137 

Callosciurus flavimanus 155 

= Sdurotamias davidianus consobrinus 308 
Thomas (1915) 19 

= Funambulus tristriatus numarius 91 

= Callosdurus 99 
Traylor, Melvin A. 9 




— Funambulus sublineatus sublineatns 95 
tristriatus, Funambulus 87 

Funambulus tristriatus 87 

Callosciurus finlaysoni 166 
Trouessart (1880) 18 

= Callosciurus flavim^nus ningpoensis 154 

= Callosciurus jlavimanus ningpoensis 154 
type description 17 
type specimen 17 

Van Gelder, R. G. 9 

= Callosciurus flavimxinvs griseimanus 148 

= Callosciurus erythraeus sladeni 113 

Tamiops svnnhoei 250 


= Callosciurus pygerythrus mearsi 222 


= Callosciurus erythraeus erythraeus 104 

Callosciurus finlaysoni 173 

= Callosciurus flavimanus styani 152 

Funambulus tristriatus 90 


= Callosciurus flavimanus rubeculus 142 

Zahn (1942) 20 

= Dremomys 259 

Callosciurus flxivimanus 136 
Zimmermann, K. 9 
Zweifel, Mrs. Richard G. 9 

Publication 991